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FLORA OF THE USSR 



Volume XXIX 

COMPOSITAE 
Tribe Cichorieae 



30 



FLORA OF THE USSR 



Initiated under the supervision and chief editorship 
of Academician V.L. Komarov 




VOL. XXIX 

COMPOSITAE 

Tribe Cichorieae 



Volume Editors 
E.G. BOBROV and N.N. TZVELEV 

General Scientific Editors 

Stanwyn G. Shetler 
Galina N. Fet 
Ellen Unumb 

Translated from Russian 




Smithsonian Institution Libraries 

Washington, D.C. 

2000 



*■ omrii 



SMIN B86 - SIL 004 

Flora SSSR, Tom. XXIX 
Akademiya Nauk SSSR Publishers, 
Moscow-Leningrad, 1964 

© 2000 Amerind Publishing Co. Pvt. Ltd., New Delhi 

Translator: Dr. B.R. Sharma 
General Editor: Dr. V.S. Kothekar 

Library of Congress Cataloging-in-Publication Data 

Flora SSSR. EngUsh 

Flora of U.S.S.R. = Flora SSSR. 

Cover title: Flora U.S.S.R. 

Translation of: Flora SSSR. 

At head of tide, v. -29. V.L. Komarov Botanical 
Institute. Academy of Sciences of the USSR. 

Reprint. Originally pubhshed: Leningrad: 
Izdatel'stvo Akademi nauk SSSR, 1934- 

Vols. Have imprint: Washington, D.C.: Smithsonian 
Institution Libraries and the National Science 
Foundation 

Includes bibliographies and indexes. 

I. Botany — Soviet Union — Classification. 
2. Plants — Identification. I. Komarov, V. L. 
(Vladimir Leont'evich), 1869-1945. II. Botanicheskii 
institut im. V.L. Komarova. III. Title: Flora SSSR. 
III. Title: Flora U.S.S.R. V. Title. 
K321.F69613 1985 85-904577 

ISBN 

Translated and published under an agreement with the Smithsonian 
Institution Libraries, Washington, D.C, by Amerind Publishing Co. Pvt. Ltd. 
66 Janpath, New Delhi 110001 

Typeset at Indira Printers, New Delhi and printed at Baba Barkha Na^h 
Printers, 26/7 Najafgar Industrial Area, New Delhi 110015. 



INSTITUTUM BOTANICUM nomine V. KOMAROVH ACADEMIAE SCIENTIARUM URSS 



FLORA 
URSS 



(FLORA UNIONIS RERUMPUBLICARUM 
SOCULISTICARUM SOVIETICARUM) 



XXK 



NAUKA 
MOSQUA - 1964 - LENINGRAD 



V.L. KOMAROV BOTANICAL INSTITUTE 
ACADEMY OF SCIENCES OF THE USSR 



Contributors 

A.G. Borissova, V.N. Vassiliev, I.T. Vassilczenko, 
M.E. Kirpicznikov, T.G. Leonova, S.J. Lipschitz, 
N.N. Tzvelev, S.K. Czerepanov, B.K. Schischkin 



SCIENTIFIC EDITOR'S PREFACE 



This is the fifth volume of six (Vols. XXV-XXX) that treat the 
large family Compositae (Asteraceae). Throughout this volume, as in 
the other volumes on this family, the translators have used the Latin 
term "capitulum" (pi., capitula), instead of the more familiar English 
term "head," for the Russian term denoting the primary inflorescence of 
florets. "Head," if it is used, denotes a secondary inflorescence, i.e., an 
arrangement of capitula. 

For practical reasons, I have concentrated my editorial review of 
this volume on the keys, discussions, and habitat and distribution 
statements. Every page of the translation has been reviewed also by 
either Dr. Fet or Ms. Unumb. Dr. Fet has paid particular attention to 
the geographic and place-name terminology in the distribution statements 
following the morphological descriptions. We are confident of the 
accuracy of the translation but also recognize that there will be 
imperfections. 

February 1998 STANWYN G. SHETLER 

Curator of Botany Emeritus 

National Museum of Natural History 

Smithsonian Institution 

Washington, DC 20560 

Assisted by Galina N. Fet 
Huntington, West Virginia 25701 

and 

Ellen Unumb 

Bethesda, Maryland. 



PREFACE 



The twenty-ninth volume contains descriptions of 595 species 
belonging to 45 genera of the subfamily Cichorioideae. The sequence 
of genera in this volume, unlike in earlier volumes of the Asteraceae 
(Compositae), corresponds to the new system proposed by George 
Ledyard Stebbins (1953), which includes some significant changes. 

This volume completes the publication of the Flora of the USSR, 
as Volume XXX was published earlier. 

Now we know that the Soviet flora comprises 17,520 species 
belonging to 1,676 genera of 160 families. 

These figures are, however, approximate. This is because inclusion 
of the plants from the Soviet Baltic Republics and the western regions 
of Belorussia, Ukraine and Moldavia in the Flora of the USSR only began 
with volumes XI-XII. Inclusion of the plants of the Kaliningrad Region, 
Karelian Isthmus and Transcarpathia, as well as southern Sakhalin, the 
Kuril Islands and Tuva, began still later, with volumes XIV-XV. 

Thus, we see that the Flora of the USSR has become outdated to a 
certain extent with the expansion of the territory of the Soviet Union. It 
has also become outdated because the publication of each volume has 
greatly facilitated the collection of new facts. 

However, only the prewar volumes published during the 1930s have 
become significantly outdated. Some of them need revision, which if 
undertaken would herald the beginning of a new series of the Flora of 
the USSR. 

Nevertheless, with the completion of the present series of the Flora 
of the USSR, we can say that now we know quite well the indigenous 
flora in general terms. 

This publication will continue to serve for the next few decades as 
a general manual for the identification of the higher plants of this country 
and as the foundation of our knowledge of the wild indigenous species. 

The fact that the territory of the Soviet Union covered by the Flora 
of the USSR constitutes one-fifth of the total land area of the earth, 
including a large part of the boreal and temperate zones of the northern 
hemisphere, imparts great international significance to this publication. 

EDITORS 



CONTENTS 



SCIENTIFIC EDITOR'S PREFACE ... vii 

PREFACE ... ix 
SYSTEMATIC INDEX OF SPECIES OF THE TWENTY-NINTH 

VOLUME OF THE "FLORA OF THE USSR" ... xiii 

SUBFAMILY II. CICHORIOIDEAE KIT AM. ... 1 

TRIBE U.Cichorieae Spreng. ... 9 

Subtribe L Scolyminae Less. ... 10 

Genus 163 L Scolymus L. ... 10 

Subtribe 2. Cichoriinae O. Hoffm. ... 13 

Genus 1632. Cichorium L. ... 14 

Genus 1633. Amoseris Gaertn. ... 19 

Genus 1634. Koelpinia Pall. ... 20 

Subtribe 3. Scorzonerinae Dum. ... 26 

Genus 1635. Scorzonera L. ... 26 

Genus 1636. Epilasia (Bge.) Benth. ... Ill 

Genus 1637. Tragopogon L. ... 115 

Genus 1638. Geropogon L. ... 197 

Subtribe 4. Hypochaerinae Less. ... 19& 

Genus 1639. Hypochaeris L. ... 199 

Genus 1640. Achyrophorus Scop. ... 201 

Genus 1641. Leontodon L. ... 204 

Genus 1642. Picris L. ... 218 

Genus 1643. Helminthia Juss. ... 227 

Genus 1644. Urospermum Scop. ... 228 

Genus 1645. Hedypnois Schreb. ... 229 

Genus 1646. Garhadiolus Jaub. and Sp. ... 230 

Genus 1647. Rhagadiolus Scop. .... 232 

Subtribe 5. Lactucinae Less. ... 234 

Genus 1648. Paramicrorhynchus Kirp. Gen. Nov. ... 235 

Genus 1649. Rhabdotheca Cass. ... 238 

Genus 1650. Atalanthus D. Don ... 240 

Genus 1651. Sonchus L. ... 243 

Genus 1652. Reichardia Roth ... 259 



Xll 



Genus 1653. Prenanthes L. 

Genus 1654. Lactuca L. 

Genus 1655. Scariola F.W. Schmidt 

Genus 1656. Steptorhamphus Bge. 

Genus 1657. Mycelis Cass. 

Genus 1658. Cephalorrhynchus Boiss. 

Genus 1659. Cicerbita Wallr. 

Genus 1660. Lagoseriopsis Kirp. Gen. Nov. 

Genus 1661. Youngia Cass. 

Genus 1662. Ixeridium (A. Gray) Tzvel. 

Genus 1663. Chorisis DC. 

Genus 1664. Paraixeris Nakai 

Genus 1665. Lapsana L. 

Genus 1666. Aposeris Neck. 

Genus 1667. Taraxacum Wigg. 

Genus 1668. Chondrilla L. 

Genus 1669. Willemetia Neck. 

Genus 1670. Heteroderis (Bge.) Boiss. 

Genus 1671. Heteracia Fisch. and Mey. 

Genus 1672. Acanthocephalus Kar. and Kir. 

Genus 1673. Crepis L. 

Genus 1674. Lagoseris M.B. 

Genus 1675. Zacintha Mill. 



261 
272 
314 
321 
331 
334 
347 
368 
370 
382 
388 
389 
395 
398 
399 
550 
577 
578 
582 
584 
585 
691 
707 



ADDENDA XXVIII. LATIN DIAGNOSES OF 
NEW SPECIES INCLUDED IN VOL. XXIX 

INDEX ALPHABETICUS 



708 
759 



VII SYSTEMATIC INDEX OF SPECIES OF THE TWENTY- 

NINTH VOLUME OF THE "FLORA OF THE USSR"* 



SUBFAMILY II. CICHORIOIDEAE KIT AM. 

Tribe 14. Cichorieae Spreng. 

Subtribe L Scolyminae Less. 

Genus 1631. Scolymus L. 

Section 1. Myscolus (Cass.) DC. 

16141. 1. S. hispanicus L. ... 12 

Section 2. Scolymus 
2. S. maculatus L. ... 13 

Subtribe 2. Cichoriinae O. Hoffm. 
Genus 1632. Cichorium L. 

1. C. intybus L. ... 16 

2. C. glandulosum Boiss. and Huet ... 18 

3. C. pumilum Jacq. ... 18 

4. C. endivia L. ... 19 

Genus 1633. Arnoseris Gaertn. 
1. A. minima (L.) Schweig. and Koerte ... 20 

Genus 1634. Koelpinia Pall. 

1. K. macrantha Winkl. ... 23 

2. K. linearis Pall. ... 24 



'Reproduced from the Russian original. The page numbers of the Russian volume 
appear in the left-hand margin of the text — General Editor. 



XIV 



16150. 3. K. turanica Vass. 

4. K. tenuissima Pavl. and Lipsch. 

Subtribe 3. Scorzonerinae Dum. 

Genus 1635. Scorzonera L. 

Subgenus I. Podospermum (DC.) Lipsch. 

1. S. lachnostegia (Woron.) Lipsch. 

2. S. laciniata L. 

3. S. calcitrapifolia Vahl. 

4. S. songorica (Kar. and Kir.) 

Lipsch. and Vass. 

5. S. armeniaca (Boiss. and Huet) 

Boiss. 

6. S. schischkinii Lipsch. and Vass. 

7. S. idae (Sosn.) Lipsch. 

8. S. grossheimii Lipsch. and Vass. 
16160. 9. S. cana (C.A.M.) O. Hoffm. 

10. S. meyeri (C. Koch) Lipsch. 
vni 11. S. kipricznikovii 'Lvpsch. 

12. S. grigoraschvilii (Sosn.) Lipsch. 

Subgenus IL Pseudopospermum (Lipsch. and Krasch.) Lipsch. 

13. S. mollis M.B. 

14. S. leptophylla (DC.) Krasch. and Lipsch. 

15. 5. suberosa C. Koch 

16. 5. turkeviczii Krasch. and Lipsch. 

17. 5. turcomanica Krasch. and Lipsch, 

18. 5. raddeana Winkl. 



25 
26 



37 
38 
39 

40 

41 

42 
42 
44 
44 
46 
47 
48 



49 
50 

52 
52 
53 
54 



Subgenus III. Scorzonera 
Section 1. Incisae Lipsch. 
16170. 19. S. bicolor Freyn and Sint. 

Section 2. Purpureae Lipsch. 

20. 5. purpurea L. 

21. 5. rosea Waldst. and Kit. 

Section 3. Egregiae (Kult.) Lipsch. 

22. S. tau-saghyz Lipsch. and Bosse 



55 



56 
58 



59 



XV 



Section 4. Piptopogon C.A.M. 

23. S. albicaulis Bge. 

24. S. bracteosa Winkl. 

25. S. alaica Lipsch. 

26. S. albertoregelia Winkl. 

27. S. tragopogonoides Rgl. and Schmalh. 

28. S. petrovii Lipsch. 
16180. 29. S. baldshuanica Lipsch. 

Section 5. Fibrillosae Nakai 

30. 5. subacaulis Lipsch. 

31. S. crispa M.B. 

32. S. ruprechtiana Lipsch. and Krasch. 

33. S. pratorum (Krasch.) Stankov 

34. 5. austriaca Willd. 

35. S. dianthoides (Lipsch. and Krasch.) 

Lipsch. 

36. S. curvata (Popl.) Lipsch. 



61 
62 
64 
64 
65 
66 
66 



67 
68 
69 
69 
70 

72 
72 



16190. 



Section 6. Pulvinares (Boiss.) Lipsch. 

37. S. filifolia Boiss. 

38. S. lipskyi Lipsch. 

39. S. seidlitzii Boiss. 

40. S. rigida Auch. ex DC. 

41. 5". koslovskyi Sosn. 



73 
74 
75 
75 
76 



Section 7. Scorzonera 



42. S. radiata Fisch. 

43. S. humilis L. 



77 
78 



IX 



16200. 



Section 8. Parviflorae Lipsch. 

44. S. parviflora Jacq. 

45. S. mongolica Maxim. 

Section 9. Turkestanicae Lipsch, 

46. S. franchetii Lipsch. 

47. S. turkestani'ca Franch. 

48. S. iliensis Krasch. 

49. S. transiliensis M. Pop. 



79 
80 



81 
82 
83 
84 



XVI 



Section 10. Foliosae (Boiss.) Lipsch. 

50. S. gracilis Lipsch. 

51. 5. stricta Homem. 

52. 5". pubescens DC. 

53. 5. hispanica L. 

54- S. inconspicua Lipsch. ex Pavl. 

55. S. chantavica Pavl. 



85 
85 
86 
87 
87 
88 



Section 11. Polycladae DC. 

56. 5. acanthoclada Franch. 

57. S. kuhistanica M. Pop. 

58. S. pseudodivaricata Lipsch. 



89 
90 
91 



Section 12. Pusillae Lipsch. 
16210. 59. S. pusilla Pall. 



92 



Section 13. Papposae Lipsch. and Krasch. 



16220. 



60. 


S. tadshikorum Krasch. and Lipsch. 


93 


61. 


S. ferganica Krasch. 


94 


62. 


S. ovata Trautv. 


95 


63. 


5. crassifolia Krasch. and Lipsch. 


95 


64. 


S. papposa DC. 

Section 14. Hissaricae Lipsch. 


96 


65. 


S. hissarica Winkl. 

Section 15. Nervosae Lipsch. 


97 


66. 


S. sosnovskyi Lipsch, 


98 


67. 


S. ketzkhovelii Sosn. 


98 


68. 


S. latifolia (Fisch. and Mey.) DC. 


99 


69. 


S. pulchra Lorn. 


100 


70. 


5. safievii Grossh. 
Section 16. Vierhapperia Lipsch. 


101 


71. 


5. ensifolia M.B. 


102 


72. 


5. biebersteinii Lipsch. 


103 


73. 


S. ramosissima DC. 


104 



XVll 



16230. 



Section 17. Tuberosae Lipsch. 
Subsection 1. Lanatae Lipsch. 

74. S. lanata (L.) Hoffm. 

75. S. pseudolanata Grossh. 

Subsection 2. Tuberiferae Lipsch. 

76. S. tuberosa Pall. 

77. S. sericeolanata (Bge.) Krasch. 

and Lipsch. 

78. S. bungei Krasch. and Lipsch. 

79. S. gageoides Boiss. 

80. S. circumflexa Krasch. and Lipsch. 

81. S. litvinovii Krasch. and Lipsch. 

82. 5. oxiana M. Pop. 

Genus 1636. Epilasia (Bge.) Benth. 

1. E. mirabilis Lipsch. 

2. E. acrolasia (Bge.) Clarke 

3. E. hemilasia (Bge.) Clarke 

Genus 1637. Tragopogon L. 
Section 1. Majores (Artemcz.) Kuthath, 



105 
105 



106 

107 
107 
108 
109 
110 
110 



112 
112 
114 



16240. 



1. 


T. dubius Scop. 


131 


2. 


T. capitatus S. Nikit. 


132 


3. 


T. paradoxus S. Nikit. 


135 


4. 


T. serawschanicus S. Nikit. 


135 


5. 


T. pseudomajor S. Nikit. 


135 



Section 2. Kemularia Boriss. 

6. T. kemulariae Kuthath. 

Section 3. Angustissimi Kuthath. 

7. T. sosnovskyi Kuthath. 

8. r. segetus Kuthath. 

9. T. vedenskyi M. Pop. ex Pavl. 

Section 4. Macropogon (Kuthath.) Boriss. 
10. T. macropogon C.A. Mey. 



136 



137 
138 
138 



139 



XVlll 



Section 5. Krascheninnikovia Boriss. 

11. T. krascheninnikovii S. Nikit. 

Section 6. Tragopogon 

12. T. orientalis L. 

13. T. pratensis L. 

16250. 14. T. bjelorussicus Artemcz. 

15. T. trachycarpus S. Nikit. 

16. T. songoricus S. Nikit. 

17. T. turkestanicus S. Nikit. 

18. T. badachschanicus Boriss. 

19. T. altaicus S. Nikit. 

20. T. karelinii S. Nikit. 

21. T. tomentosulus Boriss. 

22. T. gracilis D. Don 

23. T. subalpinus S. Nikit. 
16260. 24. T. sibiricus Ganesch. 

25. T. cretaceus S. Nikit. 



140 



142 
143 
143 
144 
145 
146 
147 
147 
148 
148 
150 
151 
151 
152 



XI 



16270. 



Section 7. Tuberosi (Kuthath.) Boriss. 

26. T. tuberosus C. Koch 

27. T. pusillus M.B, 

Section 8. Profundisulcati Kuthath. 

28. T. meskheticus Kuthath. 

29. T. ketzkhovelii Kuthath. 

30. T. plantagineus Boiss. and Huet 

31. T. buphtalmoides (DC.) Boiss. 

32. T. tasch-kala Kuthath. 

33. T. karjaginii Kuthath. 

34. T. latifolius Boiss. 

35. T. acanthocarpus Boiss. 

36. T. idae Kuthath. 

37. T. makaschwilii Kuthath. 

38. T. armeniacus Kuthath. 



152 
153 



154 
155 
155 
156 
157 
157 
158 
158 
159 
159 
160 



Section 9. Bessera Boriss. 

39. T. lithuanicus (DC.) Boriss. 

40. T. gorskianus Rchb. 

41. T. tanaiticus Artemcz. 

42. T. heterospermus Schweigg. 



162 
163 
163 
164 



XIX 



Section 10. Brevirostres Kuthath. 

43. T. dasyrhynchus Artemcz. 
16280. 44. T. volgensis S. Nikit. 

45. T. podolicus (Bess, ex DC.) Artemcz. 

46. T. elatior Stev. 

47. T. graminifolius DC. 

48. T. brevirostris DC. 

49. T. serotinus Sosn. 

50. r. filifolius Rehm. ex Boiss. 

51. T. colchicus Alb. 

52. T. stepposus (S. Nikit.) Stank. 

53. T. ucrainicus Artemcz. 

16290. 54. T. daghestanicus (Artemcz.) Kuthath. 
55. T. borystenicus Artemcz. 



165 
166 
167 
168 
169 
169 
170 
171 
171 
172 
173 
174 
174 



Section 11. Sosnovskya Boriss. 

56. T. marginatus Boiss. and Buhse 

57. T. reticulatus Boiss. and Huet 

58. T. charadzeae Kuthath. 

59. T. maturatus Boriss. 



175 
176 
177 
177 



Section 12. Dubianskya Boriss. 
60. T. subulosus Krasch. and S. Nikit. 



178 



Section 13. Dasypogon Boriss. 
61. T. kopetdaghensis Boriss. 



179 



16300. 



Section 14. Nikitinia Boriss. 

62. T. dubianskyi Krasch. and Nikit. 

63. T. scoparius S. Nikit. 

64. T. ruthenicus Bess, ex Krasch. 

65. T. donetzicus Artemcz. 



180 
182 
183 
184 



xn 



Section 15. Chromopappus Boriss. 

66. T. coloratus C.A. Mey. 

67. T. nachitschevanicus Kuthath. 



185 
185 



Section 16. Rubriflori Boiss. 
68. T. collinus DC 



186 



XX 



16310. 



69. 


r. rMfcer S.G. Gmel. 


187 


70. 


T. marginifolius Pavl. 


188 


71. 


T. conduplicatus S. Nikit. 


189 


72. 


r. montanus S. Nikit. 


190 


73. 


r. fl/flicM5 S. Nikit. 


191 


74. 


T. gaudanicus Boriss. 


191 


75. 


T. elongatus S. Nikit. 


192 


76. 


T. malikus S. Nikit. 


194 


77. 


T. kultiassovii M. Pop. 


195 


78. 


T. kasahstanicus S. Nikit. 

Section 17. Hebecarpus Boriss. 


195 


79. 


T. porrifolius L. 


196 



16320. 



Genus 1638. Geropogon L. 
1. G. hybridus (L.) Sch. Bip. 

Subtribe 4. Hypochaerinae Less. 

Genus 1639. Hypochaeris L. 
Section 1. Hypochaeris 

1. H. glabra L. 

Section 2. Porcellites (Cass.) DC. 

2. H. radicata L. 

Genus 1640. Achyrophorus Scop. 
Section 1. Achyrophorus 

1. A. maculatus (L.) Scop. 

Section 2. Oreophila (D. Don) DC. 

2. A. ciliatus (Thunb.) Sch. Bip. 

3. A. uniflorus (Vill.) Bluff, and Fingerh. 

Genus 1641. Leontodon L. 
Subgenus I. Scorzoneroides (Moench) V. Vassil. 

1. L. autumnalis L. 

2. L. pseudotaraxaci Schur. 

3. L. rilaensis Hayek 



197 



199 



200 



202 



202 
203 



207 
208 
209 



XXI 



16330. 



XIII 



4. 


L. keretinus Nyl. 


210 


5. 


L. gutzulorum V. Vassil. 


210 


6. 


L danubialis Jacq. 


211 


7. 


L. repens Schur 


211 


8. 


L. schischkinii V. Vassil. 


212 


9. 


L. croceus Haenke 

Subgenus II. Leontodon 


214 


10. 


L. asperus (Waldst. and Kit.) Boiss. 


215 


11. 


L. asperrimus (Willd.) Boiss. 


216 


12. 


L. kotschyi Boiss. 


217 


13. 


L. hispidus L. 


217 


14. 


L. caucasicus (MB.) Fisch, 


218 



16340. 



Genus 1642. Picris L. 
Section Picris 



1. p. hieracioides L. 


221 


2. P. japonica Thumb. 


221 


3. P. kamtschatica Ldb. 


222 


4. P. rigida Ldb. 


223 


5. P. similis V. Vassil. 


223 


6. P. pauciflora Willd. 


224 


7. P. strigosa M.B. 


226 


8. P. canescens (Stev.) V. Vassil. 


227 



Genus 1643. Helminthia Juss. 
1. H. echioides (L.) Gaertn. 

Genus 1644. Urospermum Scop. 
1. U. picroides (L.) Desf. 

Genus 1645. Hedypnois Schreb. 

1. H. cretica (L.) Willd. 

2. H. persica M.B. 

Genus 1646. Garhadiolus Jaub. and Sp. 

1. G. papposus Boiss. and Buhse 

2. G. angulosus Jaub. and Sp. 



228 



162 



230 
230 



231 
232 



XXll 



Genus 1647. Rhagadiolus Scop. 

16350. 1. R. edulis Gaertn. 

2. R. hebelaenus (DC.) Vass. 

Subtribe 5. Lactucinae Less. 
Genus 1648, Paramicrorhynchus Kirp. 
1. P. procumbens (Roxb.) Kirp. 

Genus 1649. Rhabdotheca Cass. 
1. R. korovinii (M. Pop.) Kirp. 

Genus 1650. Atalanthus D. Don 
1. A. acanthodes (Boiss.) Kirp. 

Genus 1651. Sonchus L. 
Section 1. Sonchus 

1. S. transcaspicus Nevski 

2. S. palustris L. 
XIV 3. S. sosnowskyi Schchian 

4. S. arvensis L. 

5. S. oleraceus L. 
16360. 6. S. asper (L.) Hill 

Genus 1652. Reichardia Roth 
1. R. dichotoma (Vahl) Freyn 

Genus 1653. Prenanthes L. 
Subgenus I. Prenanthes 

1. P. purpurea L. 

2. P. tatarinowii Maxim. 

3. P. cacaliifolia (M.B.) Beauverd 

4. P. pontica (Boiss.) Leskov 

Subgenus II. Nabalus (Cass.) Babcock, Stebbins and Jenkins 

5. P. maximowiczii Kirp. 

Species of Uncertain Affinity 

6. P. abietina (Boiss.) Kirp. 



234 
234 



237 



240 



241 



245 
247 
248 
249 
254 
256 



261 



264 
265 
266 
268 



269 



270 



XXlll 



7. P. angustifolia Boulos ... 273 

Genus 1654. Lactuca L. 

Subgenus I. Mulgedium (Cass.) Babcock, Stebbins and Jenkins 

1. L. sibirica (L.) Benth. ex Maxim. ... 280 

16370. 2. L. tartarica (L.) C.A. Mey. ... 282 

3. L. winkleri Kirp. ... 285 

Subgenus II. Lactucopsis (Sch. Bip.) Babcock, Stebbins and Jenkins 

4. L. stricta Waldst. and Kit. ... 288 

5. L. chaixii Vill. ... 289 

6. L. wilhelmsiana Fisch. and Mey. ... 290 

Subgenus III. Pterachaenium (Kitam.) Kirp. 

7. L. indica L. ... 292 

8. L. triangulata Maxim. ... 294 

9. L. raddeana Maxim. ... 295 



16380. 



Subgenus IV. Lactuca 

Section 1. Lactuca 

10. L. serriola Tomer 
ILL. sativa L. 

12. L. saligna L. 

13. L. altaica Fisch. and Mey. 

14. L. georgica Grossh. 



297 
299 
301 
303 
306 



Section 2. Micranthae Boiss. 

15. L. rosularis Boiss. 

16. L. glauciifolia Boiss. 

17. L. auriculata DC. 

18. L. undulata Ldb. 



307 
308 
309 
310 



XV 



Species of Uncertain Affinity 



19, L. spinidens Nevski 

20. L. mira Pavl. 



312 
313 



16390. 



Genus 1655. Scariola F.W. Schmidt 

1. S. viminea (L.) F.W. Schmidt 

2. S. orientalis (Boiss.) Sojak 



318 
320 



XXIV 



3. S. albertoregelia (C. Winkl.) Kirp. 

Genus 1656. Steptorhamphus Bge. 
Section 1. Steptorhamphus 



322 



1. 


S. crambifolius Bge. 


326 


2. 


S. crassicaulis (Trautv.) Kirp. 


327 


3. 


S. persicus (Boiss.) 0. and B. Fedtsch. 
Section 2. Tuberosi (Boiss.) Kirp. 


334 


4. 


S. tuberosus (Jacq.) Grossh. 


331 


5. 


S. petmeus (Fisch. and Mey.) Grossh. 


331 


6. 


S. linczevskii Kirp. 


333 


7. 


5. czerepanovii Kirp. 


334 



Genus 1657. Mycelis Cass. 

1. M. muralis (L.) Dum, 

Genus 1658. Cephalorrhynchus Boiss. 
Section 1. Cephalorrhynchus 

16400. 1. C. tuberosus (Stev.) Schchian 

2. C. talyschensis Kirp. 

3. C. kirpicznikovii Grossh. 

4. C soongoricus (Rgl.) S. Kovalevsk. 

5. C kossinskyi (Krasch.) Kirp. 

6. C. subplumosus S. Kovalevsk. 

7. C. takhtadzhianii (Sosn.) Kirp. 

Section 2. ZoUikoferiastrum Kirp. 

8. C. polycladus (Boiss.) Kirp. 

Genus 1659. Cicerbita Walk. 
Subgenus I. Cicerbita 

1. C. a/pma (L.) Walk. 

2. C. thianschanica (Rgl. and Schmalh.) 

Beauverd 
16410. 3. C. flzwrea (Ldb.) Beauverd 



336 



340 
342 
343 
346 
347 
348 
349 



350 



355 
356 

358 



XXV 

Subgenus II. Platyachena Kiq). 
Section 1. Platyachena Kirp. 

4. C. uralensis (Rouy) Beauverd ... 360 

5. C. macrophylla (Willd.) Wallr. ... 361 

6. C. sevanensis Kirp. ... 362 

7. C. racemosa (Willd.) Beauverd ... 363 

XVI Section 2. Prenanthopsis Kirp. 

8. C. prenanthoides (M.B.) Beauverd ... 365 

9. C. bourgaei (Boiss.) Beauverd ... 366 

Section 3. Multicostaria Kirp. 

10. C. deltoidea (M.B.) Beauverd ... 368 

Subgenus III, Poicilachena Kirp. 

11. C. zeravschanica M. Pop. ex ... 369 

S. Kovalevsk. 

12. C. rosea (M. Pop. and Vved.) ... 370 

S. Kovalevsk. 
16420. 13. C. kovalevskiana Kirp. ... 372 

Genus 1660. Lagoseriopsis Kirp. 
1. L. popovii (Krasch.) Kirp. ... 374 

Genus 1661. Youngia Cass. 
Section 1. Stenophytum Babe, and Stebbins 

1. Y. stenoma (Turcz.) Ldb. ... 378 

Section 2. Crepidopsis Babe, and Stebbins 

2. Y. diversifolia (Ldb.) Ldb. ... 380 

3. Y. tenuifolia (Willd.) Babe, and Stebbins ... 381 

4. Y. altaica (Babe, and Stebbins) Czer. ... 383 

5. Y. tenuicaulis (Babe, and Stebbins) Czer. ... 385 

Section 3. Pseudoixeridopsis Czer. 

6. Y. serawschanica (B. Fedtsch.) 

Babe, and Stebbins ... 386 



XXVI 



Genus 1662. Ixeridium (A. Gray) Tzvel. 

1. L. chinense (Thunb.) Tzvel. 

2. /. strigosum (Levi, and Vaniot) Tzvel. 
16430. 3. /. gramineum (Fisch.) Tzvel. 

4. /. graminifolium (Ldb.) Tzvel. 

5. /. dentatum (Thunb.) Tzvel. 

Genus 1663. Chorisis DC. 
1, C. repens (L.) DC. 

Genus 1664. Paraixeris Nakai 
Section 1. Paraixeris 

1. P. denticulata (Houtt.) Nakai 

2. P. pinnatipartita (Makino) Tzvel. 

3. P. serotina (Maxim.) Tzvel. 

Section 2. Pseudomycelis Tzvel. 

4. P. saxatilis (A. Baran.) Tzvel. 

Genus 1665. Lapsana L. 

1. L. communis L. 

2. L. intermedia M.B. 
XVII 16440. 3. L. grandiflora M.B. 

4. L. aipetriensis Vass. 

Genus 1666. Aposeris Neck. 
1. A. foetida (L.) Less. 

Genus 1667. Taraxacum Wigg. 
Section 1. Spuria (DC.) Hand.-Mazz. 

1. T. montanum (C.A.M.) DC. 

2. T. syriacum Boiss. 

3. T. serotinum (Waldst. and Kit.) Poir. 

Section 2. Taraxacum 
Subsection 1. Gymnophylla Hand.-Mazz. 

4. T. officinale Wigg. 

5. T. almaatense Schischk. 



390 
390 
391 
392 
392 



394 



397 
398 
399 



400 



401 

402 
403 
404 



404 



430 
430 
431 



433 
434 



XXVll 



6. T. complicatum S. Koval. 

7. T. ecomutum S. Koval. 
16450. 8. T. turgaicum Schischk. 

9. T. majus Schischk. 

10. T. grossheimii Schischk. 

11. T. macrolepium Schischk. 

12. T. macrochlamideum S. Koval. 

13. T. praestans Lindb. f. 

14. T. platycranum Dahlst. 



435 
435 
436 
437 
437 
438 
438 
439 
440 



16460. 



16470. 



xvm 



16480. 



Subsection 2. Alpina Schischk. 

15. T. nigricans (Kit.) Rchb. 

16. T. fontanum Hand.-Mazz, 

17. T. alpinum Hegetschw. and Heer 

Subsection 3. Spectabilia (Dahlst.) Schischk. 

18. T. platypecidum Diels 

19. T. ohwianum Kitam. 

20. T. coreanum Nakai 

21. T. mexicanum DC. 

22. T. ussuriense Kom. 

23. T. neo-sachalinense Koidz. 

24. T. tatewakii Kitam. 

25. T. natschikense Kom. 

26. T. pseudoglabrum Dahlst. 

27. T. pseudoalpinum Schischk. 

28. T. schischkinii V. Korol. 

29. T. croceum Dahlst. 

30. T. lapponicum Kihlm. ex Hand.-Mazz. 

31. T. sagittifolium Lindb. f. 

Subsection 4. Ceratophora Hand.-Mazz. 

32. T. chirieanum Kitam. 

33. T. pseudolasianthum Koidz. 

34. T. nairoense Koidz. 

35. T. grandisquamatum Koidz. 

36. T. miyakei Kitam. 

37. T. sachalinense Kitam. 

38. T. shikotanense Kitam. 

39. T. ketoiense Tatew. 

40. T. yetrofuense Kitam. 

41. 71 shimushirense Tatew. and Kitam. 



440 
441 
441 



442 
443 
444 
444 
445 
446 
446 
448 
448 
449 
449 
450 
450 
451 



452 
453 
453 
454 
454 
455 
455 
456 
456 
457 



XXVlll 



42. T. shumushuense Kitam. 

43. T. longipes Kom. 

44. T. perlatescens Dahlst. 

45. T. malaisei Dahlst. 

46. T. hultenii Dahlst. 

47. T. ceratophorum (Ldb.) DC. 
16490. 48. T. longicome Dahlst. 

49. T. acricome Dahlst. 

50. T. anchorifolium Kom. 

51. T. kljutschevskoanum Kom. 

52. T. evittatum Dahlst. 

53. T. platyceras Dahlst. 

54. T. koraginense Kom. 

55. T. yamamotoi Koidz. 

56. T. latisquameum Dahlst. 

57. T. chamissonis Greene 
16500. 58. T. kojimae Kitam. 

59. T. macilentum Dahlst. 

60. T. koraginicola Kom. 

61. T. leptoceras Dahlst. 

62. T. dilutum Dahlst. 

63. T. vulcanorum Koidz. 

64. T. hjeltii (Dahlst.) Dahlst. 

65. T. murmanicum Orlova 

66. T. norvegicum (Dahlst.) DahlsL 

67. T. simulum Brenner 
16510. 68. T. novae-zemliae Holmboe 

69. T. brevicome Dahlst. 

70. T. brachyceras Dahlst. 

71. T. confusum Schischk. 

72. T. schelkovnikovii Schischk. 

Subsection 5. Macrocomuta (Van Soest) Schischk 

73. T. glaucanthum (Ldb.) DC. 

74. T. glaucivirens Schischk. 

75. T. klokovii Litvinenko 

76. T. aldtavicum Schischk. 

77. T. contristans S. Koval. 
16520. 78. T. bicorne Dahlst. 

79. T. elongatum S. Koval. 

80. T. juzepczukii Schischk. 

81. T. botschantzevii Schischk. 

82. T. multiscaposum Schischk. 



457 
458 
459 
459 
460 
460 
461 
462 
462 
463 
463 
463 
464 
464 
465 
465 
466 
467 
467 
468 
468 
469 
469 
470 
470 
471 
471 
472 
473 
473 
474 



475 
476 
476 
477 
477 
478 
479 
479 
480 
480 



XXIX 



83. T. lipskyi Schischk. 

84. T. nikitinii Schischk. 

85. T. comitans S. Koval. 

XIX 86. T. monochlamydeum Hand.-Mazz. 

87. T. praecox Schischk. 
16530. 88. T. holophyllum Schischk, 

89. T. maracandicum S. Koval, 

90. T. strobilocephalum S. Koval. 

91. T. nuratavicum Schischk, 

92. T. nevskii Juz. 

93. T. microspermum Schischk. 

94. T. aschabadense Schischk, 

95. T. tadshicorum Ovcz, 

96. T. karatavicum Pavl, 

97. T. longipyramidatum Schischk, 
16540. 98, T. neo-lobulatum Van Soest 

99, T. butkovii S, Koval, 

100. T. kok-saghyz Rodin 

101. T. brevicomiculatum V. Korol. 

102. T. pseudo-dissimile Van Soest 

103. T. longirostre Schischk. 



481 
482 
482 
483 
483 
484 
485 
485 
486 
486 
487 
487 
488 
488 
489 
490 
490 
491 
492 
492 
493 



16550. 



Subsection 6. Obliqua (Dahlst.) Schischk. 

104. T. obliquum (Fr,) Dahlst, 

105. T. xerophilum Markl. 

106. T. dissimile Dahlst. 

107. T. stenolobum Stschegl. 

108. T. printzii Dahlst. 

109. T. desertorum Schischk. 



494 
494 
495 
496 
496 
497 



16560. 



Subsection 7. Erythrosperma (Dahlst.) Schischk. 

110. T. erythrospermum Andrz. 

111. T. pineticola Klok. 

112. T. tauricum Kotov 

113. T. intercedens Markl. 

114. T. praticola Schischk. 

115. T. armeniacum Schischk. 

116. r, androssovii Schischk. 

117. T. repandum Pavl. 

118. T. tianschanicum Pavl. 

119. T. sumneviczii Schischk. 

120. T. rubiginans Dahlst. 



498 
499 
499 
500 
501 
501 
502 
502 
503 
503 
504 



XXX 



XX 



121. 


T. 


rufum Dahlst. 


505 


122. 


T. 


lateritium Dahlst. 


505 


Subsection 8. Mongolica (Dahlst.) Kitam. 




123. 


T. 


mongolicum Hand.-Mazz. 


506 


124. 


T. 


ikonnikovii Schischk. 


507 


125. 


T. 


badachschanicum Schischk. 


508 


126. 


T. 


stanjukoviczii Schischk. 


508 


127. 


T. modestum Schischk. 


509 


16570. 128. 


T. 


saposhnikovii Schischk. 


509 


129. 


T. 


atrans Schischk. 


510 


130. 


T. 


vassilczenkoi Schischk. 


511 


131. 


T. 


kirghizicum Schischk. 


511 


132. 


T. 


songoricum Schischk. 


512 


133. 


T. 


compactum Schischk. 


512 


134. 


T. 


altaicum Schischk. 


513 


135. 


T. 


pingue Schischk. 


513 


136. 


T. 


schugnanicum Schischk. 


514 


137. 


T. 


magnum V. Korol. 


515 


16580. 138. 


T. 


eriopodum (D. Don) DC. 


515 


139. 


T. 


mbtzovii Schischk. 


515 


140. 


T. 


perpusillum Schischk. 


516 




Section 3. Palustria (Dahlst.) Schischk. 




141. 


T. 


decolorans Dahlst. 


517 


142. 


T. 


suecicum Hagl. 


517 


143. 


T. 


lissocarpum (Dahlst.) Dahlst. 


518 


144. 


T. 


balticum Dahlst. 


519 


Section 4. Laevia (Hand.-Mazz.) Schishk. 




145. 


T. 


platylepium Dahlst. 


520 


146. 


T. 


lyngeanum Hagl. 


520 


147. 


T. 


turiense Orlova 


521 


16590. 148. 


T. 


nivale Lge. ex Kihlm. 


522 


149. 


T. 


tundricola Hand.-Mazz. 


522 


150. 


T. 


glabrum DC. 


523 


151. 


T. 


atratum Schischk. 


524 


152. 


T. 


subglaciale Schischk. 


524 


153. 


T. 


arcticum (Trautv.) Dahlst. 


526 


154. 


T. 


sibiricum Dahlst. 


526 


155. 


T. 


kamtschaticum Dahlst. 


527 



XXXI 



Section 5. Porphyrantha Schischk. 

156. T. porphyranthum Boiss. 

157. T. pseudoroseum Schischk. 
16600. 158. T. lilacinum Krassan. ex Schischk. 



528 
528 
526 



Section 6. Scariosa Hand.-Mazz. 

159. T. hybernum Stev. 

160. T. pobedimoviae Schischk. 

161. T. ciscaucasicum Schischk. 



530 
531 

532 



16610. 



Section 7. Rhodotricha Hand.-Mazz. 

162. T. bessarabicum (Homem.) Hand.-Mazz. 

163. T. oliganthum Schott and Kotschy 
ex Hand.-Mazz. 

164. T. daralagesicum Schischk. 

165. T. stenolepium Hand.-Mazz. 

166. T. voronovii Schischk. 

167. T. fedtschenkoi Hand.-Mazz. 

168. T. varsobicum Schischk. 



533 

533 
534 
535 
536 
536 
537 



XXI 



Section 8. Parvula Hand.-Mazz. 

169. T. dissectum (Ldb.) Ldb. 

170. T. collinum DC. 

171. T. baicalense Schischk. 

172. T. leucanthum (Ldb.) Ldb. 

173. T. dealbatum Hand.-Mazz. 

174. T. czuense Schischk. 

175. T. sinicum Kitag. 



538 
539 
539 
540 
541 
541 
542 



Section 9. Orientale Hand.-Mazz. 

176. T. crepidiforme DC. 

177. T. heptapotamicum Schischk. 
16620. 178. T. goloskovkovii Schischk. 

179. T. tzvelevii Schischk. 

180. T. pojarkoviae Schischk. 

181. T. glabellum Schischk. 

182. T. brevirostre Hand.-Mazz. 

183. T. pseudominutilobum S. Koval. 

184. T. minutilobum M. Pop. ex S. Koval. 

185. T. erostre TLdi)^. 

186. T. rubidum Schischk. 



543 
544 
544 
545 
546 
546 
547 
547 
548 
549 
549 



XXXll 



16630. 



16640. 



187. 


T. lyratum (Ldb.) DC. 


550 


188. 


T. aksaicum Schischk. 

Species of Uncertain Affinity 


551 


189. 


T. alaicum Schischk. 


551 


190. 


T. albescens Dahlst. 


552 


191. 


T. calcareum V, Korol. 


553 


192. 


T. eriobasis S. Koval. 


554 


193. 


T. gnezdilloi S. Koval. 


554 


194. 


T. linczevskyi Schischk. 


554 


195. 


T. litwinowii Schischk. 


555 


196. 


T. luridum Hagl. 


555 


197. 


T. muricatum Schischk. 


556 


198. 


T. oschense Schischk. 


557 


199. 


T. pamiricum Schischk, 


557 


200. 


T. pseudonigricans Hand.-Mazz. 


558 


201. 


T. seravschanicum Schischk. 


558 


202. 


T. tenuisectum Somm. and Lev. 


559 



Genus 1668. Chondrilla L. 



16650. 



xxn 



Subgenus I. Chondrilla 
Section 1. Chondrilla 



1 . C. juncea L. 


565 


2. C. graminea M.B. 


566 


3. C. brevirostris Fisch. and Mey 


568 


4, C. acantholepis Boiss. 


569 


5. C. latifolia M.B. 


570 


6. C. aspera (Schrad. ex Willd.) Poir. 


571 


7. C. canescens Kar. and Kir. 


572 


8. C. gibbirostris M. Pop. 


573 


Section 2. Arthrorhynchus Fisch. and Mey, 




9. C piptocoma Fisch. and Mey. 


574 


10. C, lejosperma Kar, and Kir. 


574 


11. C, rouillieri Kar. and Kir. 


576 


12. C, laticoronata Leonova 


576 


13. C phaeocephala Rupr. 


577 


14. C. maracandica Bge. 


578 


15, C, omata Iljin 


579 



XXXlll 

Section 3. Pachyrhynchus Leonova 

16660. 16. C. kusnetzovii Iljin ... 580 

Subgenus II. Brachyrhynchus (Iljin) Leonova 

17. C. ambigua Fisch. ex Kar. and Kir. ... 580 

18. C. macrocarpa Leonova ... 581 

19. C. pauciflora Ldb. ... 582 

20. C. macra Iljin ... 584 

21. C. mujunkumensis Iljin and Igolk ... 584 

22. C bosseana Iljin ... 585 

Species of Uncertain Affinity 

-. C. hispida (Pall.) Poir. ... 586 

Genus 1669. Willemetia Neck. 

1. W. tuberosa Fisch. and Mey. ... 587 

Genus 1670. Heteroderis (Bge.) Boiss. 

1. H. pusilla (Boiss.) Boiss. ... 588 

2. H. leucocephala (Bge.) Leonova ... 590 

Genus 1671. Heteracia Fisch. and Mey. 

16670. 1. H. szovitsii Fisch. and Mey. ... 591 

Genus 1672. Acanthocephalus Kar. and Kir. 

1. A. benthamianus Rgl. ... 593 

2. A. amplexifolius Kar. and Kir. ... 594 

Genus 1673. Crepis L. 
Section 1. Hapalostephium (D. Don) Froel. 

1. C. sibirica L. ... 608 

2. C. paludosa (L.) Moench ... 610 

3. C. caucasica C.A.M. ... 612 

4. C. glabra Boiss. ... 613 

Section 2. Succisocrepis Sch. Bip. ex Bisch. 

5. C hokkaidoensis Babe. ... 615 

6. C chrysantha (Ldb.) Turcz. ... 617 

7. C. burejensis Fr. Schmidt. ... 618 



XXXIV 



16680. 8. C. polytricha (Ldb.) Turcz. 

9. C. czuensis Serg. 

10. C. karakuschensis Czer. 

Section 3. Hieracioides Froel. 

11. C. lyrata (L.) Froel. 

12. C. mollis (Jacq.) Aschers. 

13. C. willemetioides Boiss. 



620 
621 
622 



623 
624 
626 



XXIII 



16690. 



Section 4. Soyeria (Monn.) Benth. 

14. C. pontica C.A.M. 

15. C. conyzifolia (Gouan) Dalla Torre 

Section 5. Crepis 
Subsection 1. Corymbiforme (Babe.) Czer. 

16. C darvazica Krasch. 

17. C. sonchifolia (M.B.) C.A.M. 

18. C. ciliata C. Koch 

19. C biennis L. 

20. C. pannonica (Jacq.) C. Koch 

Subsection 2. Divaricatae (Babe.) Czer. 

21. C khorassanica Boiss. 



628 
629 



632 
634 
636 
637 
639 



642 



Subsection 3. Strictae (Babe.) Czer. 
22. C. turcomanica Krasch. 



643 



16700. 



Section 6. Macropodes Babe. 

23. C willdenowii Czer. 

24. C. alike ri Tamamsch. 

25. C oreades Schrenk 

26. C crocea (Lam.) Babe. 

Section 7. Ixeridopsis Babe. 

27. C. corniculata Rgl. and Schmalh. 

28. C. flexuosa (Ldb.) Clarke 

29. C alaica Krasch. 

30. C. karelinii M. Pop. and Schischk. 

ex Czer. 

31. C. sogdiana (Krasch.) Czer. 



645 
646 
647 
648 



650 
651 
653 

656 
657 



XXXV 



xxrv 



32. C lactea Lipsch. 

33. C. nana Richardson 

Section 8. Intybellioides Froel. 

34. C. praemorsa (L.) Tausch 

35. C. caucasigena Czer. 

Section 9. Mesophylion Babe. 

36. C. bungei Ldb. 

37. C. tectorum L. 
16710. 38. C. ramosissima Urv. 

39. C. astrachanica Stev. ex Czer. 

40. C nigrescens Pohle 

Section 10. Phaecasium (Cass.) DC. 

41. C. pulchra L. 

Section 11. Microcephalum Babe. 

42. C gmelinii (L.) Tausch 

43. C. multicaulis Ldb. 

Section 12. Alethocrepis Bisch. 

44. C. capillaris (L.) Walk. 

45. C. micrantha Czer. 
— . C. foliosa Babe. 



658 
659 



661 
663 



665 
667 
669 
671 
672 



674 



678 
679 



682 
684 
685 



Section 13. Barkhausia (Moench) Gaudin 

46. C. alpina L. 

47. C. foetida L. 
16720. 48. C. rhoeadifolia M.B. 

49. C. trichocephala (Krasch.) V.V. Nikit. 

50. C. kotschyana Boiss. 

Section 14. Lepidoseris (Rchb.) Babe. 

51. C. marschallii (C.A.M.) F. Schultz. 

Section 15. Nemauchenes (Cass.) Benth. 

52. C. setosa Haller f. 



688 
689 
690 
692 
693 



696 



698 



XXXVl 



Genus 1674. Lagoseris M.B. 

Subgenus I. Lagoseris 

1. L. robusta Czer. 

2. L. purpurea (Willd.) Boiss. 

3. L. callicephala Juz. 

4. L. sahendi (Boiss. and Buhse) Czer. 

Subgenus II. Pterotheca (Cass.) Czer. 

5. L. sancta (L.) K. Maly 
16730. 6. L. glaucescens (C. Koch) Sosn. 

7. L. obovata (Boiss. and Noe) Bomm. 

8. L. macrantha (Bge.) Iljin 

9. L. aralensis (Bge.) Boiss. 



704 
705 
706 
707 



708 
710 

712 
713 
714 



16735. 



Genus 1675. Zacintha Mill. 
1. Z. verrucosa Gaertn. 



716 



Volume XXX, published earlier, contains genus 1676, Hieracium 
L., with 785 species. Thus, the Flora of the USSR comprises 160 families, 
1,676 genera, and 17,520 species of vascular plants. 



Subfamily II. CICHORIOIDEAE Kitam. in Mem. Coll. Sc. Univ. 
Kyoto, ser. B, XIII (1937) 4 (Compos. Jap. I) and Ibid. XXII, 1 (1955) 
77 (Compos. Jap. IV). — CI. Syngenesia: Polygamia aequalis L. Sp. pi. 
(1753) 789, p. p. — Semiflosculosae Berkh. Expos, charact. struct. 
Compos. (1760) 6. — CI. Compositae ordo 1. Compositae semiflosculosae 
P.P. Gmel. Otia bot. (1760) 122. — Compositae sect. Lactucae Adans. 
Fam. pi. II (1763) 111.— CI. Ligulatae Hill, Hort. Kew. (1769) 39.— 
Chicoraceae Neck, in Acta Acad. Theod.-palat. II (1770) 463. — 
Cichoraceae Juss. Gen. pi. (1789) 168 (pro ord.); Link, Handb. I (1829) 
779; D. Don in Edinb. New Philos. Journ. VI (1829) 305-306 (pro 
fam.). — Cichoriaceae Juss. 1. c. nom. fam. conserv. — Glossariphytum 
Neck. Elem. bot. I (1790) 44. — Compositiflorae Ligulatae Gaertn. De 
fruct. et sem. pi. (1791) 353. — Symphianthera. Flores compositi corollulis 
omnibus ligulatis Moench, Meth. (1794) 30, 532. — CI. Compositae Fam. 
I. Semiflosculosae Batsch, Tab. affinit. regni veget. (1802) 250. — 
Chicoracees ou Semiflosculoses DC. in Ann. Mus. Hist. Nat. XVI (1810) 
145. — Lactucees ou Chicoracees Cass, in Nouv. Bull. Sci. Soc. Philom. 
Paris (1812) 189. — Compositae, Ordnung V. Cichoreae Spreng. Anleit. 
ed. 2, II (1818) 577. — Stirps Flosculacia ordo Ligulacia [fam.] 
Chicoraceae Dum. Comment, bot. (1822) 55. — Compositae A. 
Homoianthae a. Lactuceae Rchb. Consp. regni veget. (1828) 97. — CI. 
Compositae, ordo Synanthereae XX. Lactucea Bartl. Ordin. natur. pl. 
(1830) 145. — Compositae D. Homoianthae a. Cichoriaceae Rchb. Fl. 
germ. exc. (1831) 248. — Compositae subordo Liguliflorae DC. Prodr. V 
(1836) 9 and Ibid. VII, 1 (1838) 74.— Compositae C. Homoianthae a. 
Liguliflorae 1. Cichorieae Rchb. Repert. herb, sive Nomencl. (1841) 
94. — Compositae B. Liguliflorae O. Hoffm. Nat. Pflanzenfam. IV, 5 
(1893) 350. — Lactucaceae Bessey in Ann. Missouri Bot. Garden 2 (1915) 
164 (pro fam.).— Lactucoideae Solbrig in Taxon. XII, 6 (1963) 230.— 
All florets ligulate. Ligules five-toothed at apex, yellow or, less often, 
with other colors (light blue, lilac, etc.). Pollen grains predominantly 
spiny-cristate (echinulate). Plants almost always with latex contained in 
more or less welldeveloped system of anastomosing laticifers. Leaves 
alternate, only rarely opposite; often arranged in basal rosette. 

Type of subfamily: Cichorium L. 



KEY TO GENERA OF TRIBE CICHORIOIDEAE' 

1. Receptacle densely covered with coriaceous or cartilaginous- 
coriaceous floral scales, enclosing achenes and deciduous with 
latter. Capitula surrounded by spiny bracteal leaves. Strongly 
prickly annual or biennial plants of Taman peninsula, Crimea, 
and Transcaucasia 1631. Scolymus L. 

+ Receptacle without coriaceous floral scales. Capitula not 
surrounded by spiny bracteal leaves 2. 

2. All achenes, or (less often) at least some, with distinct pappus 
("letuchka") of simple or plumose hairs (bristles) 12. 

+ Achenes entirely without pappus or latter coronate (i.e. as short 
membranous, toothed, ciliate or fimbriate crown); sometimes, but 
only inner, achenes with short pappus of barbed (less often 
membranous, broadened at base) hairs (bristles) 3. 

3. All achenes without pappus as well as beak; only sometimes with 
slightly raised, scarcely toothed, and membranous frill, but then 
stems below capitula hollow, distinctly clavately swollen 4. 

+ Achenes with at least very short but quite distinct pappus 7. 

4. Achenes 8-15 mm long. Inner involucral bracts (much longer than 
outer ones) enlarged on fruiting, hardened and enclosing peripheral 
stellate-flattened achenes 1647. Rhagadiolus Scop. 

+ Achenes shorter, mostly up to 5 mm long. Inner involucral bracts 
not enlarged on fruiting 5. 

5. Plants with more or less numerous small capitula on slender 
peduncles, forming cymose or paniculate general inflorescences. 
Achenes with numerous (about 20) fine ribs 

1665. Lapsana L. 

+ Plants scapose bearing solitary terminal capitula, or stems slightly 
broadened, but then latter clavately swollen below capitulum...6. 

6. Perennial, with pinnatipartite or pectinately partite rosulate leaves. 
Achenes 4-5 mm long, indistinctly tetra- or pentaquetrous 

1666. Aposeris Neck. 

+ Annual, with more or less distinctly sinuate-toothed, oblong- 

obovate, rosulate leaves. Achenes 1.5-2 mm long 

1633. Arnoseries Gaertn. 

7(3) Inner involucral bracts with fruits thickened and hardened, 

sometimes compactly fused with each other 8. 

+ Inner involucral bracts remaining unchanged with fruits, herbaceous 

(if so, sometimes connate at base) or thin-coriaceous 10. 



'Treatment by M.E. Kirpicznikov. 



8. Inner involucral bracts with fruits connate into hard shell-Hke cover, 
open on top (only apices free and foliaceous). Capitula pyriform- 
globose, small 1672. Acanthocephalus Kar. and Kir. 

+ Inner involucral bracts with fruits not connate 9. 

9. All achenes without beak, peripheral with coronate and toothed 
pappus, inner ones with pappus of membranous bristles. Ligulate 
florets exceeding involucres. Plants of the Caucasus, less often in 
Crimea 1645. Hedypnois Schreb. 

+ All or some achenes with beak or at least apically produced; pappus 
of short bristles or coronate — fimbriate or fimbriate-ciliate. Ligulate 
florets shghtly longer than involucres. Plants of Soviet Central Asia 

and the Caucasus 

1646. Garhadiolus Jaub. and Sp. 

10(7) Ligules mostly light blue, blue or bluish-pink, sometimes (in 
albinos) whitish. Receptacles with short split scales. Involucre more 
or less cyhndrical. Pappus coronate, very short (only 0.2-0.8 mm 
long), of small-toothed scales 1632. Cichorium L. 

+ Ligules yellow or light yellow (turning pale or sometimes blue on 
drying). Receptacle glabrous or with few bristles. Capitula with 

fruits hemispherical. Annuals 11. 

11. Basal leaves more or less crenate-dentate to pinnatifid and 
pinnatipartite, oblong or oblong-obovate; cauline leaves sessile, 
amplexicaul or semiamplexicaul, with sagittate base. Outer achenes 

persistent, inner deciduous 

1671. Heteracia Fisch. and Mey. 

+ All leaves entire, mostly linear-filiform or linear, less often 
narrowly oblong or lanceolate, not amplexicaul. All achenes 

deciduous 1634. Koelpinia Pall. 

12(2) Inner involucral bracts with fruits indurate and swollen in form of 
hood, compactly enclosing outer achenes by their margins; latter 
with beak, inner achenes without beak. Annuals with yellow 
ligules; in the USSR found only in Crimea... 1675. Zacintha Mill. 

+ Inner involucral bracts with fruits not indurate and not swollen.. 
13. 

13. All or at least one row of pappus hairs (bristles) plumose 14. 

+ Pappus hairs simple or somewhat barbed 23. 

14. Involucre one-rowed 15. 

4- Involucre two-rowed or imbricate 17. 

15. Beak at base (at articulation with achene) hollow, swollen, attached 
to achene at an angle. Pappus of one row of easily detaching white 
plumose hairs, as long as beak. Involucre of eight foliaceous bracts, 
coronate below. Annual, with yellow ligules 

1644. Urospermum Scop. 



+ Beak, if present, not swollen at base. All other characters different 
16. 

16, All achenes with pappus of similar hairs. Receptacle without scales. 
Perennials or biennials 1637. Tragopogon L. 

+ Pappus of achenes dissimilar; in peripheral achenes with five 
protruding, scabrous, persistent bristles, pappus hairs soft in inner 

achenes. Receptacle fimbriate- villous. Annual 

1638. Geropogon L. 

17(14) Receptacle with long scarious scales, exceeding achenes 18. 

+ Receptacle glabrous, less often pubescent or with short scaly or 
fimbriate scales 19. 

18. Pappus two-rowed: hairs of outer row short, almost filiform; hairs 
of inner row long, plumose. Scarious scales on receptacle not 
overlapping. Growing in European part of the USSR (excluding 
Crimea) and the Caucasus 1639. Hypochaeris L. 

+ Pappus one-rowed. Scales on receptacle arranged longitudinally. 
Perennials with rosette of rather large basal leaves, found in the 
European part of the USSR (excluding Crimea), Caucasus, and 

Siberia, less often in the Far East 1640. Achyrophorus Scop. 

19(17) Pappus hairs entangled (plumose parts of adjacent hairs entwined), 
soft 20. 

+ Pappus hairs free, not entangled 21. 

20 Achenes with callous annulus at apex or near midpoint; in latter 
case, upper part of achene covered with woolly hairs merging into 
pappus. Annuals 1636. Epilasia Bge. 

+ Achenes without callous annulus, glabrous or pubescent, at base 
sometimes with hollow appendage (pedicel; if so, leaves usually 

pinnatisect). Perennials 1635. Scorzonera L. 

21(19) Outer involucral bracts three to five (six), relatively broad 
(resembling small leaves in form) and inner row of narrowly linear, 
long, cuneately acuminate bracts (usually eight). Achenes with 
fragile filiform beak, patelliform above, bearing persistent one- 
rowed pappus. Annual weed, with stem, leaves, and capitula 
covered with stiff hairs. Found in Crimea and Caucasus (and in 
Turkmenia? — introduced) 1643. Helminthia Juss. 

+ Outer involucral bracts absent. Combination of other characters 

different 22. 

22. Pappus two-rowed: inner row of basally fused plumose hairs. Stem 
branched, leafy, mostly without rosette of basal leaves (with 
exception of Picris rigida Ldb.) 1642. Picris L. 

+ Pappus one- or two-rowed, its hairs not fused at base. Stem mostly 
scapose or weakly branched, with one or two or several small linear 
leaves. Lower leaves in basal rosette 1641. Leontodon L. 



23(13) Receptacle with less numerous setiform scales much longer than 

achenes 1674. Lagoseris M.B. 

+ Receptacle mostly glabrous, smooth to pitted or alveolate, with 
scaly margins; very rarely short ciliate or fimbriate, but always 
without setiform scales 24. 

24. Stem leafless (scapose), almost always hollow (only in very rare 
cases stem weakly branched, and then capitula two or several), 
terminating in single apical capitulum. Perennial with basal leaves 
arranged in rosette 25. 

+ Stem leafy, almost always more or less strongly branched; capitula 
very rarely one or several, usually more or less numerous, in 
corymbose, corymbose-paniculate, paniculate (less often racemose 
or other type) general inflorescence 26. 

25. Involucre mostly two-rowed (less often incompletely three-rowed). 
Plants with slender creeping rhizome, at places tuberous. Achenes 
with 2-2.5 mm long flattened part and beak about four times as 
long, with crown of five (six) short, apically roundish scales at 
conically broadened base. Within the USSR, growing only in 
Caucasus 1669. Willemetia Neck. 

+ Involucre of several rows of bracts. Plants mostly with cord-like 
root. Combination of remaining characters different. Genus with 

species distributed throughout the USSR 

1667. Taraxacum Wigg. 

26(24) Achenes heterogenous: outer achenes few, subtended by inner 

involucral bracts, without or with very short beak and easily 

detaching pappus; inner achenes numerous, prismatic, apex sharply 

) tuberculate, with beak and persistent pappus, beak two times as 

long as body of achene. Annual herbs with few-flowered capitula 

and rosette of basal leaves. Found in Soviet Central Asia 

1670. Heteroderis (Bge.) Boiss. 

+ Achenes all alike; if different, then inner and outer achenes 

differing from each other in various characters 27. 

27. Body of all or only inner achenes tuberculate or with various types 
of scales along ribs, especially at apex (or simultaneously with 
tubercles and scales). Beak often articulated, breaking easily at 
articulation with pappus, or without articulation, but then almost 
always clavately thickened at apex. Sometimes achenes smooth or 
almost so, but then with crown of five short scales at apex, and 
distinctly articulated beak. Very rarely even crown absent (or 
almost indistinct), but then involucral bracts and flowers in 
capitulum five each, and plant with thick, flexuous stem, often 

divaricately branched. Perennials, less often biennials 

1668. ChondrUla L. 



+ Achenes without tubercles and scales; combination of renaaining 
characters different 28. 

28. Completely glabrous herb with prostrate branches, rooting at nodes; 
lamina divided into three to five almost palmate segments. Achenes 
with rather thick porous pericarp. Perennials. Found in the Far East 

1663. Chorisis DC. 

+ Plants more or less pubescent; if glabrous, not rooting at nodes. 
Remaining characters also different 29. 

29. Achenes heteromorphic, at least peripheral ones appreciably 
different from inner ones in form, nature of pubescence, or in other 
characters; occasionally some achenes with pappus, and others 
without it 30. 

+ Achenes homomorphic, i.e., all achenes in capitulum practically 
similar 36. 

30. Inner hairs of pappus relatively thick and long, less numerous (five 
to nine); outer hairs more numerous, short, soft and silky. Achenes 
10-12 mm long, narrowly cylindrical (columnar). Plants endemic 
to Soviet Central Asia 1649. Rhabdotheca Cass. 

+ Pappus (sometimes present only in some achenes) hairs similar in 
thickness. Achenes smaller or of different form 31. 

3 1 . Outer achenes prismatic or columnar, tetra- or pentaquetrous, dark 
brown, with four to five longitudinal furrows, transversely 
tuberculate. Flowers in capitulum (35)50-70(100). Perennials, 
growing in the Caucasus 1652. Reichardia Roth 

+ Outer achenes of different form. Flowers in capitulum almost 
always not many 32. 

32. Highly branched perennial or semishrub with hard, prickly, terminal 
branches. Leaves dropping off early, older plants becoming leafless, 
virgate. Capitula 5-6(1 0)-flowered, solitary or (less often) from two 
to a few, mostly on secondary branches. Endemic to Turkmenia 

1650. Atalanthus D. Don 

+ Annual or perennial herbs without prickly terminal branches. 
Remaining characters different 33. 

33. Pappus about three times as long as achenes, of very fine and silky 
white hairs connate at base into small ring. Outer achenes with 
rather numerous prominent, longitudinal, tuberculate-rugose ribs, 
narrowed toward apex, and terminating in very small, readily 
detaching beak. Plants growing in Soviet Central Asia 

1643. Paramicrorhynchus Kirp 

+ Pappus shorter than achene, as long or sometimes slightly longer. 
Remaining characters different 34. 

34. Capitula few-flowered, only four to six flowers in each. Outer 
achenes with uncinate outgrowths, densely covered, as also entire 



achene, with very short distant hairs (seen under hand lens!); inner 
achenes more or less glabrous. Annual, endemic to deserts of Soviet 
Central Asia 1660. Lagoseriopsis Kirp. 

+ Capitula usually many-flowered 35. 

35. Corolla tube 2-3 mm long, densely villose. Inner achenes 3.5-5(6) 
mm long 1673. Crepis L. [sect. Phaecasium (Cass.) DC] 

+ Corolla tube 4-6(8) mm long, with scattered short, needle-shaped 
hairs or almost glabrous. Inner achenes (including beak) 9-18 mm 

long 1673. Crepis L. [sect. Barkhausia (Moench) Gaudin] 

36(29) Involucre of only one row of well developed bracts. Capitula few- 
flowered, each with three to five yellow flowers. Herbs with short 
rhizome, growing in the European part of the USSR and Caucasus 
1657. Mycelis Cass. 

-H Involucre of two or more distinct rows of bracts; if one-rowed, 
flowers in capitulum at least eight 37. 

37. Pappus of outer crown of short, light colored hairs (seen under hand 
lens!), and inner row of well developed hairs; sometimes outer 

I crown indistinct but then achenes flattened, (1.5)2-3 mm wide, 

with beak, or achenes slightly flattened, with small dark spots (seen 

under hand lens!) also on narrowed apex 38. 

-I- Outer crown of short hairs absent; all pappus hairs more or less 
similar; achenes almost always narrower, or without beak and 
purple spots 40. 

38. Achenes with well-developed slender beak 39. 

+ Achenes without beak; their apex with either short, wide, somewhat 

deep constriction, or wide neck 1659. Cicerbita Wallr. 

39. Achenes (1.5)2-3 mm wide, slightly to strongly flattened, with one 
or few (mostly up to three) prominent longitudinal ribs 

1656. Steptorhamphus Bge. 

+ Achenes up to 1 mm wide, fusiform, with more or less numerous 
prominent longitudinal ribs. Beak differing in color from body of 

achene. 1658. Cephalorrhynchus Boiss. 

40(37) Cauline leaves long, linear, decurrent on intemodes and attached 
to stem by their bases (auricles) in form of prominent green stripes 
on lighter-colored stem. Capitula few-flowered, mostly with five 
flowers each 1655. Scariola F.W. Schmidt 

+ Cauline leaves attached to stem without bases 41. 

41. Lower leaves narrowed into winged petiole, with broad scarious 
base, along margin and inward mostly covered with papillate and 
stellate hairs. Plants endemic to Darvaz 

1655. Scariola F.W. Schmidt 

+ Lower leaves different 42. 

42. Achenes strongly flattened, almost black, elliptical or oblong- 



elliptical, in broadest part (1.5)2-2.5 mm wide, with one or a few 
prominent longitudinal ribs, beak very short or absent, apex with 
wide collar. Biennial or perennial, mostly with underground 
narrow-fusiform thickenings, growing almost exclusively in the Far 

East, less often in Eastern Siberia '. 

1654. Lactuca L. (subgenus Pterachenium (Kitam.) Kirp.) 

+ Achenes not or only slightly flattened or differently colored and 
narrower; remaining characters different 43. 

43. Achenes without beak or with beak-like tip, only sometimes 
slightly narrowed toward apex 44. 

+ Achenes with more or less distinct beak, usually differing in 
structure and color from body, (beak sometimes very short — 
visible under hand lens!), or achenes narrowed into beak-like 
tip, mostly pale in color, less often lighter 49. 

44. Ligules light blue, blue or lilac; capitula with 15-20(25) flowers 
each. Leaves lanceolate or oblong-lanceolate. Weeds or 

semiweeds 1654. Lactuca 

L. (subgenus Mulgedium (Cass.) Babe, Stebb. and Jenk.). 

+ Ligules yellow; if light blue-lilac, then number of flowers in 
capitulum or shape of leaves different 45. 

45. Fully developed capitula with 75 or more flowers each; ligules 
yellow; achenes somewhat to strongly flattened, ovoid, oblong, 
obovoid, or less often more or less prismatic. Pappus hairs smooth 
or almost so 165L Sonchus L. 

+ Flowers in each capitulum fewer and if with same number, 
achenes or pappus different 46. 

46. Capitula with 9-15 flowers each and yellow ligules, often turning 
pink after flowering. Achenes somewhat flattened, with ribs of 
different thickness. Outer as well as inner involucral bracts with 
distinct, narrow, bent cornicle on outer side 

1661. Youngia Cass. 

-I- Capitula with larger number of flowers, if flower as many or 
less, their ligules colored otherwise or achenes of different shape 
47. 

47. Achenes fusiform, narrowed toward apex, with 10 or more 
longitudinal ribs. Pappus of white or whitish hairs. Capitula many- 
flowered; ligules almost always yellow 1653. Prenanthes L. 

+ Achenes not narrowed toward apex, cylindrical, prismatic, oblong- 
linear or linear , 48. 

48. Achenes oblong-linear or linear, somewhat flattened or less often 
almost cylindrical, with faint striations or with more or less 
distinct longitudinal four to five, less often more ribs. Pappus of 
numerous dirty brown or light rust-colored, less often white hairs. 



Flowers in each capitulum 5, 8-10, 15 or 25-35 

1653. Prenanthes L. 

+ Achenes cylindrical or prismatic, truncate (and not narrowed as 
in Crepis; cf. couplet 47), mostly with 10 longitudinal ribs. Pappus 
almost always of stiff, bristly, yellowish or dirty white hairs, 
extreniely rarely of white flexible hairs. Flowers in capitulum 

mostly numerous, very rarely 20-45 Hieracium L.' 

49(43) Beak or beak-like tip shorter than body of achene 50. 

10 + Beak or beak-like tip more or less as long as body of achene, 

often much longer. Plants usually more or less pubescent 

52. 

50. Plants glabrous, growing in the Far East, and less often also in 
eastern Siberia 51. 

+ Plants pubescent. Annuals, growing in the European part of the 

USSR (including Crimea) and in Caucasus 

1673. Crepis L. (sect. Nemauchenses (Cass.) Benth.) 

51. Pappus hairs persistent, up to 3.5-7.0 mm long. Perennials, usually 
with persistent basal leaves and a few cauline leaves 

1662. Ixeridium (A. Gray) Tzvel. 

+ Pappus hairs readily detaching; achenes 2.5-3.5 mm long. Annuals 
and biennials with early-withering basal leaves, but with numerous 

cauline leaves 1664. Paraixeris Nakai 

52(49) Capitula many-flowered (with more than 25 flowers each). 
Achenes fusiform, narrowed toward base, with 10 longitudinal 
puberulent ribs (seen under high magnification). Pubescent 
biennials with woody vertical root, found in Crimea and Caucasus. 

1673. Crepis L. (sect. Lepidoseris (Rchb.) Babe). 

+ Capitula (8)10-20(25)-flowered. Achenes more or less distinctly 
to strongly flattened, narrow-obovoid or oblong-ellipsoidal, with 
rather numerous or less often only one to three longitudinal ribs; 
achenes puberulent along ribs (seen under hand lens!), and 
between them (seen under high magnification), usually in 
transverse flexuous rows 

1654. Lactuca L. (subgenus Lactuca). 

Tribe 14. CICHORIEAE Spreng. Syst. Veget. Ill (1826) 633 
("Cichoreae"; pro tribu e cl. Syngenesiae). — Lactuceae Cass.' in Bull. 



'The descriptions of Hieracium L. species occupy a separate volume published earlier. 
Cf. the Flora of the USSR, Vol. 30, Izd. Akad. Nauk SSSR, Moscow-Leningrad, 1960. 



10 

Soc. Philom. (1815) 173, in Journ. Phys. (1819) 151 (non vidi) and in 
Diet. sc. nat. XX (1821) 355, XXV (1822) 59, XLVIII (1827) 422 (pro 
tribu Synantherearum); Dum. Fl. Belg. (1827) 59. — Trib. Cichorieae 
Dum. Analyse fam. pi. (1829). — Cichoraceae Less, in Linnaea V (1830) 
132 and in Synops. Compos. (1832) 126 (pro tribu Synantherearum); 
11 DC. Prodr. VII, 1 (1838) 74 (pro tribu Compositarum).— Nexus 
Compositae 1. Synanthereae, trib. Liguliflorae Griseb. Grundr. system. 
Bot. (1854) 136. — Trib. Cichoriaceae (Juss.) Benth. in Benth. and Hook, 
f. Gen. pi. II (1873) 168, 219.— [Trib.] Cichorieae Baill. Hist. pi. VIII 
(1882) 70, 105.— Trib. Cichorieae Rchb. ex O. Hoffm. in Engl, and 
Prantl. Nat. Pflanzenfam. IV, 5 (1893) 350.— The description of the 
tribe corresponds to the characterization of the subfamily. 

Type of tribe: type of subfamily. 

Subtribe 1. SCOLYMINAE Less. Synops. Compos. (1832) 126 
("Scolymeae"); DC. Prodr. VII, 1 (1838) 75; Benth. in Benth. and Hook, 
f. Gen. pi. II (1873) 168, 219; O. Hoffm. in Engl, and Prantl. Nat. 
Pflanzenfam. IV, 5 (1893) 353; Stebbins in Madrono XII, 3 (1953) 70.— 
Trib. Lactuceae sect. 1. Archetypae Cass, in Diet. sc. nat. XXV (1822) 
60, p. min. p. — Trib. Lactucees sect. 1. Prototypes 1. Scolymees Cass, 
in Diet. sc. nat. XLVIII (1827) 422.— Trib. Cichoreae D. Don in Edinb. 
New Philos. Journ. VI (1829) 307. p. p. (solum gen. Scolymus). — Trib. 
Cichorieae g) Scolymeae Dum. Analyse fam. pi. (1829) 30. — Pappus 
coronate, only sometimes corona with a few long scabrous bristles. Pollen 
grains spiny-cristate (echinulate). Receptacle with coriaceous or 
cartilaginous-coriaceous scales enclosing achene from sides and falling 
with it. Basic chromosome number: jc = 10. 

Type of subtribe: Scolymus L. 



GENUS 1631. Scolymus L.'^ 

L. Sp. pi. (1753) Sl3.—Myscolus Cass, in Bull. Soc. Philom. (1818) 33. 
Capitula usually rather numerous, less often (in very small 



'Classifications of the Lactuceae were published twice by A. Cassini: first in 1822 (in 
Diet. sc. nat. XXV, pp. 59-89), when a brilliant treatment of this group of the Compositae 
[Asteraceae] was given after the list of genera; in 1827, A. Cassini (in Diet. sc. nat. XLVIII, 
pp. 421-435) slightly modified and supplemented the classification of the Lactuceae. The 
latest publications of his classification (in Diet. sc. nat. LX, 1830, pp. 568-569; in Opusc. 
phytol. Ill, 1834, pp. 44 4 6) repeated the classification published in 1827 and, therefore, 
are not cited by us. 



11 

specimens) solitary, terminal on stems and lateral branches, and sessile 
or almost sessile in axils of upper cauline leaves, enclosed by two to 
six spiny terminal leaves, homogamous, with rather numerous (15-40) 
bisexual ligulate florets. Involucre ovate, 12-16 mm long and 4-7 mm 
wide, consisting of a few irregular rows of imbricate, coriaceous- 
herbaceous, lanceolate or lanceolate-linear bracts, gradually increasing 
in length from outer to inner. Receptacle obtusely conical, densely 
covered with ovate (from oblong-ovate outer to broadly ovate inner) 
coriaceous or cartilaginous-coriaceous, 4-8 mm long scales; bracts on 
inner (ventral) side with two longitudinal folds, enclosing ovary and 
achene on sides and falling with achene. Flowers yellow or light yellow, 
usually almost one and one-half times as long as involucre; corolla 12- 
12 25 mm long, with four or five rather long teeth at apex and more or 
less hairy tube. Anthers sagittate at base, and with rounded triangular 
appendage at apex. Style branches relatively short, almost filiform. 
Achenes 2-4 mm long, oblong, strongly flattened dorsally, with five 
longitudinal ribs, of which two lateral ribs very prominent, and ventral 
and two dorsal ones significantly weak, glabrous, tip with short (0.2- 
0.7 mm long) irregularly toothed crown and two long and scabrous 
barbed bristles from its base, or only with crown without bristles; achenes 
on all sides (with exception of their tip with crown) firmly enclosed by 
bracts and their longitudinal folds, falling together with them. Annual 
or biennial herbs with more or less branched erect stems and more or 
less crenate-dentate or pinnatipartite highly spiny, sessile, decurrent 
cauline leaves. 

Type of genus: S. maculatus L. 

Three species of the genus are represented in the Mediterranean 
Region, from the Pyrenees and North Africa to the Caucasus. Of these, 
two species are found in the USSR. 

1 Capitula mostly in axils of middle and upper cauline leaves and 
usually surrounded by three leaves (one sheathing and two lateral); 
leaf margin not or scarcely thickened; achenes with crown and 
two long bristles at tip 1. S. hispanicus L. 

+ Capitula predominantly terminal on stem and lateral branches, 
usually enclosed by four to six terminal leaves; leaf margin with 



'Treatment by N.N. Tzvelev. 

^Latin transcription of the Greek name of the plant — skolymos, originating from the 
word skolos — spine or thorn. 



12 

moniliform thickening; achenes with only crown at tip, without 
bristles 2. S. maculatus L. 

Section 1. Myscolus (Cass.) DC. Prodr. VII (1838) 76.— Myscolus 
Cass, in Bull. Soc. Philom. (1818) 33 and in Diet. sc. nat. XXIV (1825) 
83. — Achenes with crown and two long bristles at tip; capitula 
predominantly axillary. 

Type of section: S. hispanicus L. 

1. S. hispanicus L. Sp. pi. (1753) 813; M.B. Fl. taur.-cauc. II, 262; 
DC Prodr. VII, 76; Ldb. Fl. Ross. II, 770; Boiss. R. on III, 713; Schmalh. 
Fl. II, 132; Grossh. Fl. Kavk. IV (1934) 223 and Opredel. Rast. Kavk. 
(1949) 502; Stank, and Tal. Operedel. Vyssh. Rast. (1949) 681; Kuthath. 
in Fl. Gruzii VIII (1952) 595. — Myscolus microcephalus Cass, in Diet. 
sc. nat. XXXIV (1825) 85.— M. hispanicus (L.) Dietr. Synops. pi. IV 
(1839-1852) 1333.— Ic: Rchb. Ic. Fl. Germ. XIX (1858) t. 1352; Jav. 
and Csap. Ic. Fl. Hung. (1933) 551. 

Annual or biennial. Strongly spiny, 15-60 cm high plant, covered 
with rather sparse, arachnoid, flexuous hairs and scattered spinules. Stem 
13 highly branched almost from base, erect. Basal leaves early- withering; 
cauline leaves numerous, oblong-lanceolate to oblong-ovate, more or less 
pinnatipartite or crenate-dentate, with lobes and teeth terminating in stiff 
spines, margin not or weakly thickened, base decurrent on stem in form 
of spiny wings, usually slightly narrowed below. Capitula usually rather 
numerous, in axils of middle and upper cauline leaves as well as terminal 
on branches, usually surrounded by three leaves (one sheathing and two 
lateral). Involucre 12-16 mm long; receptacular scales relatively thin 
and coriaceous. Flowers yellow. Achenes 2-4 mm long with short (0.3- 
0.7 mm long) crown and two long scabrous bristles at tip. July to October 

Sands and gravel-beds at seacoast and in river valleys, often as a 
weed of the roadsides and inhabited places. — European Part: Lower 
Don (Taman Peninsula), Crimea (southern part); Caucasus: Eastern 
Transcaucasia (Muganskaya steppe). Western Transcaucasia (near 
seacoast). General distribution: Southern Europe (north to Hungary and 
Romania), northern Africa, West Asia, Canary Islands, Described from 
southern Europe (southern France, Italy). Type in London. 

Section 2. Scolymus — Scolymus sect. Euscolymus DC. Prodr. VII 
(1838) 75. — Achenes with only crown at tip, without bristles; capitula 
predominantly terminal. 

Type of section: type of genus. 

2. S. maculatus L. Sp. pi. (1753) 813; DC. Prodr. VII, 75; Boiss. 
Fl. or. Ill, 713; Schmalh. Fl. II, 132; Stank, and Tal. Opredel. Vyssh. 
Rast. (1949) 681.— Ic: Rchb. Ic. R Germ. XIX (1858) t. 1353. 



13 

Annual or biennial. Strongly spiny, 15-60 cm high plant, covered 
with very soft flexuous hairs (to almost glabrous) and rather numerous 
spinules. Stem branched almost from base, erect. Basal leaves early- 
withering; cauline leaves numerous, oblong-lanceolate to lanceolate-ovate, 
pinnatifid or pinnatipartite, with lobes terminating in very strong long 
spines, margin strongly thickened (in form of whitish ridge), base long 
decurrent on stem in form of spiny wings usually not narrowed below. 
Capitula usually rather numerous, terminal on stem and lateral branches, 
less often in axils of upper cauline leaves, usually enclosed by four to 
six spiny terminal leaves. Involucre 12-16 mm long; receptacular scales 
cartilaginous-coriaceous. Flowers yellow. Achenes 2.5-4 mm long, with 
very short (0.2-0.3 mm long) and narrow crown at tip, without bristles. 
Flowering July to October. 

Only as an introduced weed of the roadsides and inhabited places. — 
European Part: Crimea (reported by Schmalhausen on the basis of a 
14 single specimen from the herbarium of W.G. Besser without precise 
indication of locality). General distribution: southern Europe, northern 
Africa, Asia Minor, Canary Islands. Described from southern Europe 
(Italy and southern France). Type in London. 

Subtribe 2. CICHORIINAE O. Hoffm. in Engl, and Prantl. Nat. 
Pflanzenfam. IV, 5 (1893) 353, p. min. p.; Stebbins in Madrono XII, 3 
(1953) 70. — ^Trib. Lactuceae sect. 2. Crepideae Cass, in Diet. sc. nat. 
XXV (1822) 61, p. min. p. (solum gen. Koelpinia); sect. 3. Hieracieae 
Cass. op. cit. 63, p. min. p. (solum gen. Amoseris); sect. 4. Scorzonereae 
Cass. op. cit. 64, p. min. p. (solum gen Cichorium): — Trib. Lactucees, 
sect 2. Crepidees I. Lampsanees Cass, in Diet. sc. nat. XL VIII (1827) 
422, p. min. p. (gen. Koelpinia); sect. 3. Hieraciees Cass. Ibid., p. min. 
p. (gen. Amoseris); sect. 4. Scorzonerees IV. Catanancees Cass. op. 
cit.— Subtrib. Scorzonereae Dum. Fl. Belg. (1827) 63 p. min. p. — Trib. 
Cichoreae a) Cichoreae Dum. Analyse fam. pi. (1829) 30. — Trib. 
Cichoreae D. Don. in Edinb. New Philos. Joum. VI (1829) 307, p. p. 
(solum gen. Cichorium). — Trib. Catanancheae D. Don Ibid. — Subtrib. 
Lampsaneae Less. Synops. Compos. (1832) 126, p. min. p. (solum gen. 
Koelpinia); DC. Prodr. VII, 1 (1838) 76 (eodem ambitu).— Subtrib. 
Hyoserideae Less. op. cit. 127, p. min. p.; DC. Prodr. VII, 1 (1838) 78 
(eodem ambitu); Benth. in Benth. and Hook. f. Gen. pi. II (1873) 168, 
219 p. min. p. — Subtrib. Rhagadioleae Benth. in Benth. and Hook. f. 
Gen. pi. II (1873) 168, 220, p. min. p. (solum gen. Koelpinia). — Pappus 
of very short and broad scales or pappus coronate, short-toothed; 
sometimes totally absent. Pollen grains spiny-cristate (echinulate). 
Receptacle glabrous or with less numerous bristles or scales. Basic 
chromosome number in most genera: x = 9. 

Type of subtribe: type of tribe. 



14 

GENUS 1632. Cichorium L.'^ 

L. Sp. pi. (1753) 813. 

Capitula usually rather numerous, less often (in very small 
specimens) solitary, in axils of middle and upper cauline leaves or 
terminal on stem and its lateral branches, homogamous, with rather many 
(8-20) bisexual ligulate florets. Involucre more or less cylindrical, 8-14 
mm long and 4-7 mm wide; outer involucral bracts four to seven, 
irregularly arranged or almost in one row, ovate to broadly lanceolate, 
highly unequal in shape and size (and then shorter than inner bracts), or 
almost similar (and then almost as long as inner bracts), more or less 
pubescent of glabrous, more or less coriaceous in lower half, upper half 
15 herbaceous; inner bracts five to eight, in one row or almost so, lanceolate 
or linear-lanceolate, thin coriaceous or coriaceous-herbaceous, glabrous 
or more or less hairy. Receptacle almost flat, irregularly alveolate, 
covered with very short more or less toothed or fimbriate scales along 
margins — margins of alveoles. Flowers light blue, blue, bluish-pink or 
whitish (in albinos), usually longer than involucre; corolla 10-25 mm 
long, with (four) five (seven) rather long lanceolate teeth, more or less 
hairy (especially in upper part of tube). Anthers sagittate at base, with 
obtuse roundish, triangular appendage at apex. Style branches thin and 
rather long. Achenes 2-3.5 mm long, obovoid, or oblong, outer achenes 
distinctly flattened dorsally and adnate with inner involucral bracts, 
glabrous and smooth (less often with solitary spinules in upper part), 
with indistinct veins and three to five more or less prominent ribs, apex 
truncate, with narrow and shallow constriction below crown; pappus 
very short, 0.2-0.8 mm long, of numerous more or less scaly broad 
hairs in two or three irregular rows. Perennial, biennial, or annual herbs 
with more or less branched and relatively loosely leafy erect stems. 
Leaves runcinately pinnatipartite to more or less toothed or undivided; 
basal leaves on short- winged petioles, usually in rosettes; cauline leaves 
sessile, amplexicaul. 

Type of genus: C. intybus L. 

About 10-12 species of the genus are mainly distributed in the 
region of the ancient Mediterranean from the Pyrenees and northern 
Africa to the western provinces of China; one species (C. intybus L.) is 
distributed almost throughout Europe and over a significant part of Asia; 



'Treatment by N.N. Tzvelev. 

-Latin transcription of the Greek plant name — kichorion, which itself originates from 
the words kio — to move, and chorion — field (i.e., a plant often entering the borders of 
fields). 



15 

also found as an introduced plant in many temperate and subtropical 
countries of both hemispheres. Of these, four species are found in the 
USSR. 

1. Whole plant, except involucral bracts, glabrous, less often nearly 
so (with isolated hairs in lower part of stem and along midrib of 
leaves), 30-120 cm high cultivated plant; involucral bracts more 
or less ciliate along margin, but without glands, outer bracts ovate- 
lanceolate to ovate, more or less distant, usually two-thirds as 
long as inner bracts, less often almost as long; pappus 0.4-0.8 
mm long 4. C. endivia L. 

-I- Plants more or less pubescent or setose, at least in lower part of 
stem or leaves beneath 2. 

2. Perennials (but usually short-lived); outer involucral bracts half 
to two-thirds as long as inner bracts, more or less squarrose, 
usually of unequal length and shape; corolla 15-25 mm long; 
pappus 0.2-0.3 mm long 1. C. intybus L. 

+ Annuals or biennials; outer involucral bracts more or less as long 

16 as inner bracts and appressed to them, usually almost similar in 

length and shape; corolla 9-15 mm long; pappus 0.4-0.8 mm 

long 3. 

3. Steni branches with divergent glandular hairs up to apex (bases 
of capitula); outer involucral bracts covered with numerous long 
glandular hairs in upper (fohaceous) half on both sides 

2. C. glandulosum Boiss. and Huet 

-I- Stem branches glabrous or sparsely hairy with rather long, 
flexuous, eglandular hairs or bristles at apex; outer involucral 
bracts ciliate along margin in upper half, glabrous or with 
scattered simple hairs outside, usually glabrous within 

3. C. pumilum Jacq. 

Series 1. Intybus Tzvel. — Perennial plants with relatively thick, often 
many-headed cord-like root; outer involucral bracts half to two-thirds as 
long as inner bracts, more or less divergent; corolla 15-25 nmi long; 
pappus 0.2-0.3 mm long. 

1. C. intybus L. Sp. pi. (1753) 813; M.B. Fl. taur.-cauc. II, 262; 
DC. Prodr. VII, 84, excl. p; Ldb. Fl. Ross. II, 774; Boiss. Fl. or. Ill, 
715; Schmalh. Fl. II, 134; Grossh. Fl. Kavk. IV, 223 and Opred. Rast. 
Kavk. (1949) 502; Kryl. Fl. Zap. Sib. XI, 2966; Stank, and Tal. Opred. 
Vyssh. Rast. (1949) 682; Kuthath. in Fl. Gruzii VIII (1952) 596; 
Mikhailovsk. in Fl. Beloruss. V. (1959) 159; V. Nikit. in Fl. Turkm. 
VII (1960) 285; Karjag. in Fl. Azerb. VIII (1961) 482; Kovalevskaja in 
fl. Uzb. VI (1962) 417;— Ic: Rchb. Ic. Fl. Germ. XIX (1858) t. 1357; 



16 

Syreistsch. 111. Fl. Mosk. Gub. Ill (1910) 319; Fedtsch. and Fler. Fl. 
Evrop. Ross. (1910) 1035; Majevsk. Fl. (1918) 345; Hegi, 111. Fl. VI, 2 
(1929) t. 273, fig. 4; Jav. and Csap. Ic. Fl. Hung. (1933) 551; Stank, 
and Tal. op. cit. fig. 380; Zemlinskii, Lekarstv. Rast. SSSR (1951) 291; 
Majevsk.: Fl. Ed. 8 (1945) Fig. 233; Mikhailovsk. op. cit. Plate 52.— 
Exs.: Fl. exs. Reipubl. Bohem. Sloven. No. 972. 

Perennials. Plants 20-150 cm high, green or glaucous-green; stem 
erect, usually more or less branched, branches often strongly divergent 
and slightly thickened toward apex, more or less setose or with crisped 
hairs, often glabrous or almost so. Leaves dorsally setose or crisped 
hairy, ventrally more or less crisped hairy; basal leaves usually persisting 
during flowering, runcinately pinnatipartite to undivided, but more or 
less toothed, gradually narrowed into winged petiole; cauline leaves 
(except lowermost; these similar to basal leaves) relatively less numerous 
and highly reduced, lanceolate-ovate to lanceolate, amplexicaul, with 
roundish or sagittate, often more or less toothed auricles. Capitula usually 
numerous, solitary or a few grouped together, terminal on stem and 
branches, and also in axils of middle and upper cauUne leaves. Involucre 
17 8-14 mm long; outer involucral bracts half to two-thirds as long as 
inner ones and usually more or less divergent, unequal in shape and 
length, relatively thin coriaceous in lower part, more or less ciliate along 
margin, more or less hairy on outer side, with simple or glandular hairs, 
often entirely glabrous; inner bracts usually more or less hairy or ciliate 
in upper part. Corolla 15-25 mm long, with varying shades of blue, 
sometimes whitish, rapidly discolored in water. Achenes 2-3 mm long, 
with 0.2-0.3 mm long pappus. Flowering July to October. 

Meadows, forest glades, herb slopes, often as weed near roads, in 
fields, near inhabited places; in mountains up to middle belt. — European 
Part: all regions, but in north (Karelia-Lapland, Dvina-Pechora) only as 
a rare introduced plant; Caucasus: All regions; Western Siberia: Ob' 
(lower part). Upper Tobol, Irtysh (rarely); Eastern Siberia: Angara-Sayans 
and Dauria (only as rare introduced plant); Soviet Central Asia: All 
regions. General distribution: Almost all of Europe, northern Africa, 
West Asia, Iran, western provinces of China; as introduced plant in 
South Africa, North and South America, Australia and New Zealand. 
Described from Europe. Type in London. 

Note. A more southern and xerophilous race predominates in Soviet 
Central Asia and southern and eastern Transcaucasia. — C intybus subsp. 
glaucum (Hoffmagg. and Link) Tzvel. comb, nova [= C glaucum 
Hoffmagg. and Link, Fl. Portug. II (1820) 178, t. 95], which is widely 
distributed almost throughout the Mediterranean Region. Specimens 
approaching it are also present on the southern coast of Crimea. This 
subspecies differs from typical C. intybus (subsp. intybus) by the glaucous- 



17 

green color of the entire plant, generally more divaricately branched 
stem, and entirely glabrous (without cilia and hairs) outer involucral 
bracts (inner bracts also eciliate, but pubescent at apex) and often was 
identified earlier as C divaricatum [C.A.M. Verzeichn. Pfl. Cauc. (1831) 
62] or as C. intybus var. divaricatum [Ldb. Fl. Ross. II, 774). The range 
of both subspecies of C. intybus (subsp. intybus and subsp. glaucum) 
greatly overlap, and forms intermediate between them are found quite 
frequently. In my opinion, this overlapping of the range is to a great 
extent secondary and associated with the rapid spread of both subspecies 
(especially that of subsp. intybus) under human influence in recent times. 
Economic Importance. The roots of the cultivated forms of this 
species (a whole series of varieties is known, for example, Borisovskii, 
Isopolinskii, and so on) are widely used for preparing a substitute for 
coffee ("chicory") and can be used for deriving sugar and alcohol. The 
leaves can be used as salad. The roots of the wild form are used in 
medicine for extracting a bitter substance that stimulates the activity of 
the digestive organs. A good honey-producing plant. 

Series 2. Pumila Tzvel. — Annual or biennial plants with slender 

cord-like root; outer involucral bracts usually almost as long as inner 

18 ones and appressed to them, less often (in C. endivia) shorter; corolla 

usually 9-15 mm long, less often (in C endivia) up to 23 mm long; 

pappus 0.4-0.8 mm long. 

2. C. glandulosum Boiss. and Heut in Boiss. Diagn. pi. or. ser. 2, 
III (1856) 87; Boiss. Fl. or. Ill, 716; Grossh. Fl. Kavk. IV (1934) 223 
and Opred. Rast. Kavk. (1949) 502, p. p. 

Annual or biennial. Plants 20-70 cm high, more or less glacuous- 
green. Stem erect, usually more or less branched, with branches rather 
strongly thickened toward apex, lower part glabrous or almost so, upper 
part densely covered with numerous, rather long, divergent, glandular 
hairs. Leaves usually more or less covered with curly and glandular 
hairs on both sides; basal and lower cauline leaves oblong, gradually 
narrowed into winged petiole, more or less toothed, often withering 
before flowering; cauline leaves relatively less numerous, sessile; middle 
leaves more or less narrowed and semiamplexicaul, upper leaves 
amplexicaul with roundish auricles, sharply toothed with teeth usually 
produced into long cusps. Capitula usually rather numerous, solitary, 
terminal on stem and branches. Involucre 8-12 mm long; outer involucral 
bracts almost as long as inner ones and appressed to them, coriaceous 
in lower half, herbaceous in upper half, covered with numerous rather 
long glandular hairs on both sides. Corolla 9-15 mm long, light blue, 
less often whitish. Achenes 2.5-3.5 mm long with 0.6-0.8 mm long 
pappus. Flowering June to September. 



18 

Gypsum-containing clayey slopes and screes. — Caucasus: Southern 
Transcaucasia (western parts: in the Araks River Basin). General 
distribution: Armenia and Kurdistan. Described from northeastern Turkey. 
Type in Geneva. 

3. C. pumilum Jacq. Observ. bot. IV (1771) 3, t. 80.— C. 
divaricatum Schousb. Vextr. Marok. (1800) 197; Boiss. Fl. or. Ill, 716.— 
C. intybus (3. divaricatum (Schousb.) DC. Prodr. VII, 84. — C. endivia p. 
pumilum (Jacq.) Vis. Fl. Dalm. II (1847) 97. — C. endivia subsp. pumilum 
(Jacq.) Hegi, 111. Fl. VI, 2 (1929) 998.-5. glandulosum auct. non Boiss. 
and Huet: Grossh. Fl. Kavk. IV, 223 and Opred. Rast. Kavk. (1949) 
502, p. p.; Karjag. in Fl. Azerb. VIII (1960) 483.— Ic: Jacq. op. cit.; 
Rchb. Ic. Fl. Germ. XIX (1858) t. 1357.— Exs.: Fl. exs. austro-hung. 
No. 208. 

Annual or biennial. Plants 15-50 cm high, green, often with 
glaucescent tinge. Stems erect, usually more or less branched, with 
divaricate branches strongly thickened toward apex, setose-hairy to 
completely glabrous below, glabrous or sparsely hairy (with rather long 
simple hairs or birstles) above. Leaves on both surfaces more or less 
covered with curly hairs and bristles; basal and lower cauline leaves on 
short-winged petioles, runcinately pinnatipartite to entire, more or less 
19 toothed; cauline leaves relatively few and strongly reduced, sessile, 
amplexicaul, often with more or less toothed auricles. Capitula usually 
rather numerous, solitary, terminal on main stem and branches, sometimes 
also in axils of upper cauline leaves. Involucre 8-12 mm long; outer 
involucral bracts almost as long as inner ones and appressed to them, 
coriaceous in lower half, herbaceous in upper half, long-ciliate along 
margin (but without glands), usually with a few-simple hairs on back, 
glabrous or almost so within. Corolla 9-15 mm long, light blue (?). 
Achenes 2.5-3 mm long, with 0.4-0.7 nun long pappus. Flowering July to 
October. 

Clayey and stony slopes, also frequently a weed of the roadsides, 
inhabited places, and borders of fields; found up to lower mountain 
zone. — Caucasus: Southern and Eastern Transcaucasia, Talysh. General 
distribution: southern Europe, northern Africa, West Asia, Iran; 
introduced into northern Europe. Described from northern part of Balkan 
Peninsula. Type in Vienna. 

4. C. endivia L. Sp. pi. (1950) 813; Schmalh. Fl. II, 134; Stank, 
and Tal. Opredel. Vyssh. Rast. (1949) 682.— C. endivia (3. sativa DC. 
Prodr. VII (1838) 84.-5. endivia subsp. endivia Hegi, 111. Fl. VI, 2 
(1929) 998.— Ic: Rchb. Ic, Fl. Germ. XIX (1858) t. 1358; Syreistsch. 
111. Fl. Mosk. Gub. Ill (1910) 319; Hegi, op. cit. fig. 687 and 688; Jav. 
and Csap. Ic. Fl. Hung. (1933) 551. 



19 

Annual or biennial. Plants glabrous, less often almost so, 30-120 
cm high. Stem erect, usually divaricately branched, glabrous, less often 
with isolated bristles in lower part; branches usually distinctly thickened 
toward apex. Leaves glabrous, less often with isolated bristles or curly 
hairs along midrib, relatively thin and soft, green, less often slightly 
glaucescent; basal and lower cauline leaves on short-winged petioles, 
pinnatisect to undivided, more or less toothed, often crimped; middle 
and upper cauline leaves fewer, greatly reduced, oblong to broadly ovate, 
sessile, amplexicaul, with acute or more or less obtuse auricles. Capitula 
usually numerous, solitary or several grouped together at apices of main 
stem and branches, also in axils of middle and upper cauline leaves. 
Involucre 9-14 mm long; outer involucral bracts more or less ciliate 
along margin, usually two-thirds as long as inner bracts, less often almost 
as long, coriaceous in lower part; corolla 12-33 nrni long, hght blue, 
less often whitish, not decolorized in water, Achenes 2.5-3.5 mm long, 
with 0.4-0.8 mm long pappus. Flowering July to October. 

Cultivated as salad plant in more southern areas of the USSR. — 
General distribution: Unknown in wild state: cultivated mostly in 
Mediterranean countries. Possibly described from southern Europe. Type 
in London. 

Note. C. pumilum Jacq. is usually considered to be the ancestor of 
this species, although it is not found in the wild state. However, a whole 
20 series of characters is common to C. endivia L. and C. intybus L. (for 
example, a larger corolla than in C. pumilum, shorter outer involucral 
bracts, capitula not only terminal on branches but also in groups of a 
few in the axils of cauline leaves), which suggests, as a more probable 
origin, that C endivia is the result of the hybridization of C. intybus x 
C. pumilum in the past. 

Economic Importance. Cultivated widely in Mediterranean countries 
as a salad plant; known since ancient times. There are many varieties 
differing mainly in leaf shape and color. 



GENUS 1633. Arnoseris Gaertn.'^ 

De Fruct. et Semin. pi. II (1791) 355; O. Hoffm. in Pflanzenfam. IV, 5 
(1894) 360. 

Capitula solitary or in groups of twos and threes on long peduncles, 
clavately thickened toward apex. Involucre two-rowed; inner involucral 
bracts enlarged and hardened on fruiting, enclosing peripheral achenes. 



'Treatment by I.T. Vassilczenko. 

^From the Greek words amos — lamb; and series — salad. 



20 

Receptacle glabrous. Corolla yellow. Achenes oblong-obovoid, 
pentaquetrous, and slightly compressed, small, without pappus and beak. 
Annual herbs with rosette of basal leaves and leafless flowering stems 
(scapes). 

A monotypic genus. 

1. A. minima (L.) Schweig. and Koerte, Fl. Erlang. II (1811) 72; 
Hegi, 111. Fl. VI, 2, 1004; Klokov. in Vizn. Rosl. USSR, 584; Geideman, 
Opred. Rast. Moldav. SSR (1954) 297.— A. pusilla Gaertn. De Fruct. et 
Sem. II (1791) 355; DC. Prodr. VII, 78; Ldb. Fl. Ross. II, 2, 773.— A. 
clavata Bub. Fl. Pyren. II (1900) 49.— Hyoseris minima L. Sp. pi. (1753) 
809. — Lampsana minima Lam. Encycl. meth. Ill (1791-1792) 414. — 
Ic: Hegi, op. cit. fig. 693, 694; Rchb. Ic. Fl. Germ. XIX (1858) t. 
1354.— Exs.: Fl. polon. exs. No. 361. 

Annuals. Stems 5-30 cm high, simple or weakly branched, numerous, 
usually reddish, leafless, glabrous below, clavately thickened and hollow 
above (below capitula); leaves in basal rosette, oblong-obovate, more or 
less crenate-dentate, puberulent on lower side and along margin. Capitula 
solitary terminal on main stems and branches, 5-8(10) mm wide. 
Involucral bracts oblong-lanceolate (outer bracts small and narrow, linear- 
lanceolate), somewhat fluffy, finely ciliate along margin. Ligulate florets 
22 pale to golden-yellow, one and one-half times as long as involucre. 
Achenes 1.5-2 mm long, obpyramidal, somewhat glossy, greenish-gray 
with five thick and lighter main ridges and five thinner longitudinal ribs 
in between, with barely raised, indistinctly toothed, membranous border 
at upper (broadened) end. Flowering April to September. (Plate I, Fig. 3.) 

Sandy places and light (sandy loam) soils sometimes a weed in 
fields. — European Part: Baltic Region, Bessarabia, Upper Dniester, Upper 
Dnieper. General distribution: Central Europe, Atlantic Europe, Mediter- 
ranean Region; introduced into North America, Australia, and New 
Zealand. Described from western Europe. Type in London. 

Note. Ledebour (1. c.) reported the presence of this plant near Penza 
and Moscow, which, however, has not been confirmed subsequently. 



GENUS 1634. Koelpinia Pall.' ^ 

Pall. Reise III (1776) Anhang, 755. 

Capitula small, 5-12-flowered. Receptacle glabrous or with less 
numerous bristles. Involucre one- or two-rowed; inner involucral bracts 
five to seven, much longer than outer (latter two); involucral bracts 



'Treatment by I.T. Vassilczenko. 

^A.R. Koelpin, physician and naturalist, a contemporary of P.S. Pallas. 



21 




21 Plate I. 

1 — Hedypnois cretica Willd.; 2 — Aposeris foetida (L.) Cass.; 3 — Amoseris minima (L.) 

Schweig. and Koerte. 



22 

spreading at fruiting, vertical during flowering. Flowers yellow, becoming 
pale on drying, sometimes turning blue. Achenes free, stellately 
spreading, linear-cylindrical, often falcate, with hooked spines or tubercles 
on dorsal surface, and with tuft of divergent hooked bristles at apex. 
Annual herbs with linear, filiform or oblong leaves. 
Types of genus: K. linearis Pall. 

The genus includes four or five species, distributed in western and 
Soviet Central Asia, entering only a part of Europe (in the Precaspian 
semidesert). 

1. Capitula large; flowers bright yellow, 15-20 mm long; receptacle 
with sparse hairs; achenes strongly curved (almost annular) 

1. K. macrantha Winkl. 

+ Capitula smaller; flowers pale yellow, 7-9 mm long, barely 
exceeding involucre; receptacle glabrous; achenes falcate to almost 
straight 2. 

2. Achenes slightly falcate to straight, coarse (thickened), gradually 
but slightly narrowed toward apex, 15-25 mm long, dorsally with 
relatively sparse spinules-bristles, or with rows of short spines, 
or tubercles. Growing on sands and stony slopes 

3. K. turanica Vass. 

23 + Combination of characters otherwise 3. 

3. Involucre (during flowering) 4-5 mm long; leaves linear, 1-2(5- 
7) mm wide; achenes dorsally more or less densely and uniformly 
spiny 2. K. linearis Pall. 

+ Involucre 2-2.5 mm long during flowering; leaves piliform- 
filiform, 0.5-1.0 mm wide; achenes smooth or sparsely spinulose 

(or with tubercles on spine) 

4, K. tenuissima Pavl. and Lipsch. 

1. K. macrantha Winkl. in Tr. Peterb. Bot. Sada XI, 5 (1890) 285; 
Pavlov, Fl. Tsentr. Kazakhst. Ill, 312; M. Popov in Tr. Uzb. Cos. Univ. 
Ser. Biol. 14 (1941) 88; Kovalevskaja in Fl. Uzb. VI (1962) 421.—/:. 
linearis Pall. var. macrantha (Winkl.) O. and B. Fedtsch. in Perech. 
Rast. Turk. 4 (1911) 306. — K. maracandica M. Pop. op. cit. 

Annual. Stem branched, erect or ascending, glabrous or slightly 
pubescent, 10-35 cm high. Leaves narrowly linear, linear or broadly 
linear to oblong, 8-12(15) cm long, (0.6) 1-3 (4-7) mm wide, gradually 
narrowed into petiole, one- to three-veined. Capitula one or two, on long 
peduncles; receptacle with a few (mostly three to five) bristles. Inner 
involucral bracts 10-13 mm long, linear-lanceolate, ciliate along margin, 
arachnoid hairy (sometimes, besides white, black and shorter hairs also 
present); outer bracts less numerous, almost half as long as inner. 



23 

pubescent. Flowers about 15-20 mm long, (1)1.5-2.5(3) mm wide, much 
longer than involucre, yellow, turning slightly blue on drying. Achenes 
strongly curved (usually annular), cylindrical, with numerous short hooked 
spinules on outer side, in longitudinal rows, inner side without spines, 
grooved. Flowering March to May. 

Stony, sandy and clayey slopes and plains in foothill deserts and in 
foothills, sometimes a weed in fields, orchards and on roadsides. — Soviet 
Central Asia: mountainous Turkmenia, Kara-Kum, Amu-Darya, Pamiro- 
Alai (southern). General distribution: Apparently found in adjacent 
regions of Iran and Afghanistan. Described from southern Tadzhikistan 
(Vakhsh-Kafimigan). Type in Leningrad. 

Note. The northern boundary of the distribution of this species passes 
along the Krasnovodsk-Chardzhou-Samarkand line. Popov (op. cit.) 
proposed recognition of a new species, K. marcandica M. Pop. Study of 
the material showed that both species are connected by a whole series 
of gradual transitions (with respect to length of lingules and width of 
leaves) and are also found in the same region. Therefore, there is in- 
sufficient reason to accept K. maracandica M. Pop. as a separate species. 
But within the limits of K. macrantha, there is considerable variation, 
apparently based on differences in the ecological conditions under which 
this species grows. On clayey compact soils, K. macrantha has narrow 
24 (almost filiform) leaves, and narrow ligules, not exceeding 15 mm in 
length. These plants, in particular, were included by Popov under the 
new species K. maracandica. However, plants of this type were described 
even earlier by Winkler as K. macrantha. Thus, K. maracandica M. 
Pop. should be referred to typical K. macrantha. On sands, the flowers 
of K. macrantha are larger and reach up to 15-20 mm with the ligule 
width of 2-3 mm, and the leaves also become wider (up to 2-5-7 mm 
wide); these plants can be referred to a separate variety (var. arenaria 
m.). Lasdy, the flowers of K. macrantha on stony soils become similar 
to the flowers of the preceding variety, but the leaves are narrow: 
narrowly linear to filiform (var. hyssarica m.). All these varieties are 
not very sharply distinguishable from each other and are connected by 
transitional forms. Finally, mention must be made of the discovery by 
N.A. Androsov near Samarkand of the giant form of K. macrantha with 
wide oblong-obovate leaves, as well as the discovery by Radde in 1886 
on sands near Imam-baba Station, of a pubescent form {K. macrantha 
var. raddeana Winkl. op. cit 148), 2.5-3 cm high with stiff leaves. In 
the latter case, a single, specimen was collected which also apparently 
was damaged (eaten on by animals), making it impossible to determine 
the taxonomic position of this form, although Winkler made this very 
significant notation on the label attached to this plant: "An potius nova 
species?" 



24 

2. K. linearis Pall. Reise III (1776) Anhang, 755; DC. Prodr. VII, 
78; Ldb. Fl. Ross. II, 2, 772; Jaub. and Spach. 111. pi. or. Ill (1847- 
1850) 123; Boiss. Fl. or. Ill, 721; Winkl. in Tr. Peterb. Bot. Sada, XI, 
2, 147; O. and B. Fedtschenko, Perech. Rast. Turk. 4, 306; Krasch. in 
Fl. Yugo-Vost. VI (1936) 437; Grossh. Fl. Kavk. IV (1934) 224; Pavlov, 
Fl. Tsentr. Kazakhst. Ill (1938) 332; M. Popov in Tr. Uzb. GoS. Univ. 
Ser. Biol. 14 (1941) 88; Kovalevskaja in Fl. Uzb. VI (1962) 420.— K. 
latifolia Winkl. in Tr. Peterb. Bot. Sada, XI, 5 (1890) 2%A.—K. linearis 
f. latifolia (Winkl.) O. Hoffm. in O. Paulsen, PI. Coll. As. Med. (1903) 
137.— Rhagadiolus koelpinia Willd. Sp. pi. Ill (1803) 1626.— Ic: Pall, 
op. cit. t. L, 1, fig. 2; Jaub. and Spach, op. cit., t. 286.— Exs.: GRF No. 
1884. 

Annual. Stems branched from base, glabrous or more or less 
pubescent, 5(10)-30(40) cm high. Leaves linear, oblong, up to 5-10(15) 
cm long and from 1-2 mm (with single longitudinal vein) to 5-7 mm 
wide (with three veins — var. latifolia (Winkl.) O. Hoffm.), more abundant 
in lower part of plant. Capitula one (or two) on slender axillary 
peduncles; one capitulum usually on peduncle originating from plant 
base (in middle of bush). Receptacle glabrous. Involucral bracts 
lanceolate-linear, more or less pubescent or almost glabrous; outer bracts 
two of three, short; inner bracts five to seven, longer (up to 5-8 mm). 
Flowers pale yellow, sUghtly longer than involucre. Achenes (8)12-15(16) 
mm long, light brown or light reddish-brown, usually six to eight, 
25 stellately spreading, linear-cylindrical, and falcately incurved (sometimes 
almost annular); longitudinally grooved on inner side, with numerous 
hooked spinules on outer side, gradually narrowed above with apical 
crown of ovate, divergent, bent spinules. Achenes usually finely 
pubescent or glabrous. Flowering April to May. 

Semideserts (plains and mountains), deserts (in mountains entering 
even middle zone), clayey, clayey-gravelly and stony soils; often a weed 
in irrigated fields (in oases) and unirrigated lands (in lower hills). — 
European Part: Lower Volga; Caucasus: Eastern and Southern 
Transcaucasia, Talysh; Soviet Central Asia: all regions. General 
distribution: West Asia (up to Himalayas), northern Africa. Described 
from Caspian plain (from Mt. Bogdo). Type in Leningrad. 

Note. In spite of all the attempts to maintain the species K. latifolia 
Winkl., I must disagree with this, even though K. latifolia was recognized 
as a species by such authorities as M.G. Popov, N.V. Pavlov and others. 
The main character by which K. linearis is differentiated from K. 
latifolia — leaf width (leaves narrow, 1-2 mm wide, with a single vein 
in the former species and leaves wide, 5-7 mm wide, three-veined in 
the latter species) — is not constant, and all possible transitions are found; 
sometimes both wide and narrow leaves are found even on the same 



25 

plant. As regards the presence of gray tomentose pubescence on young 
stems and involucres of K. latifolia (to which M.G. Popov drew attention, 
op. cit.), it must be mentioned that even this character is extremely 
variable and inconsistent, and, incidentally, the young involucres and 
stems of the type specimen of K. latifolia, preserved in the Herbarium 
of the Botanical Institute, Academy of Sciences of the USSR, are only 
weakly hairy, nearly glabrous. If we also consider that K. latifolia does 
not have a separate range and is found scattered over the entire range 
of K. linearis, it becomes apparent that K. latifolia is nothing more than 
a variety of the latter species and the one associated with the better 
moisture conditions. 

3. K. turanica Vass. in Flora Uzb. VI (1962) 479, 515. 
Annual, stem rough, branched, ascending, up to 20-30 cm high, 

weakly hairy. Leaves linear (or narrowly linear), 1-1.5 to 2.5-3 mm 
wide, one- veined. Capitula on rather long (1 cm or longer) peduncles, 
axillary (one usually in middle) on short, lanate peduncle. Involucral 
bracts linear-lanceolate, more or less lanate, outer bracts one-third to 
half as long as innear ones, latter reaching length of 5-8(10) mm. Dry 
ligules light colored (whitish), only slightly longer than involucres. 
Achenes slightly curved to almost straight (or even straight), coarse 
(thickened), gradually but slightly narrowed toward apex, 15-25 mm 
long, grooved along length, puberulent or glabrous, with tuft of stellate, 
26 divergent, hooked bristles at tip, and sparse bristles (uncate at apex) on 
outer side or with a row of short spinules or tubercles. Flowering March 
to April (May). 

Sands, stony slopes. — Soviet Central Asia: Aralo-Caspian, Balkhash 
Region, Kara-Kum, Kyzyl-Kum, Syr-Darya, Amu-Darya. Endemic. 
Described from Repetek. Type in Leningrad. 

4. K. tenuissima Pavl. and Lipsch. in Pavlov, Fl. Tsentr. Kazakhst. 
Ill (1938) 332; Kovalevskaja in Fl. Uzb. VI (1962) 478.—/:. leiocarpa 
M. Pop. in Tr. Uzb. Gos. Univ. Ser. Biol. 14 (1941) 88. 

Annual. Stems slender, prostrate or ascending, glabrous or sparsely 
hairy, highly branched, 5-10(15) cm high. Leaves filiform, 0.5-1 mm 
wide. Capitula less numerous, axillary, 2-3 mm wide, on rather long 
slender peduncles. Inner involucral bracts linear-lanceolate, 2.5-4(5) mm 
long, more or less hairy; outer bracts half as long. Ligules barely 
exceeding involucre, pale (cream-colored) when dry. Achenes 
glabrous or with sparse scattered spinules along dorsal surface, weakly 
curved to almost straight, with apical tuft of hooked hairs, short (7-9- 
12 mm long) and slender, four of five, gradually broadened. Flowering 
April to May. 



26 

Saline, clayey and stony deserts, outcrops of gypsum-containing clay 
and red sandstone, often on takyrs. — Soviet Central Asia: all regions. 
General distribution: Possibly found in Iran and Afghanistan. Described 
from the desert near Kzyl-Orda. Type in Moscow; isotype in Leningrad. 

Note. Pavlov (op. cit.) noted that this plant is "found in highly 
saline, clayey and stony, desert steppes" in the region of Irgiz and Turgai. 
According to Pavlov, this species is distributed over the deserts of 
southern Kazakhstan and possibly into Iran and Afghanistan. "The plant 
is observed with difficulty," wrote Pavlov, "for it grows in extreme 
desert conditions and also has extremely fast and early vegetative 
growth." 

Subtribe 3. SCORZONERINAE Dum. Fl. Belg. (1827) 63 
("Scorzonereae") p. p.; Less. Synops. Composit. (1832) 131, p. p.; DC. 
Prodr. VII, 1 (1838) 99, p. p.; Benth. in Benth. and Hook. f. Gen. pi. II 
(1873) 169, 222 p. p.; Stebbins in Madrono XII, 3 (1953) 75.— Trib. 
Lactuceae sect. 4. Scorzonereae Cass, in Diet. sc. nat. XXV (1822) 64, 
p. min. p. — Trib. Lactucees sect. 4. Scorzonerees II. Scorzonerees vraies 
Cass, in Diet. sc. nat. XLVIII (1827) 422, p.p. — Trib. Scorzonereae D. 
Don in Edinb. New Philos. Joum. VI (1829) 307.— Trib. Cichorieae b) 
Scorzonereae Dum. Analyse fam. pi. (1829) 30, p. p. — Sippe 
Tragopogoneae Sch. Bip.' in Flora XVII, 2 (1834) 476. — Leontodontinae 
O. Hoffm. in Engl. u. Prantl. Nat. Pflanzenfam. IV, 5 (1893) 361, p. min. 
p. — Pappus mostly of plumose hairs (bristles). Pollengrains spiny-cristate. 
27 Receptacle glabrous, rarely more or less pubescent. Basic chromosome 
number x = 6 and 7. Principal veins usually arcuate-parallel. Plants 
glabrous or pubescent; pubescence of soft tomentose hairs or of different 
kind. 

Type of subtribe: Scorzonera L. 



GENUS 1635. Scorzonera L 



I. 2 



L. Sp. pi. (1753) 790. 

Capitula more or less large, cylindrical or obconical. Involucre 
oblong, almost cylindrical; involucral bracts many-rowed, imbricate, 
gradually increasing in size from outside inward, without (subgenus 
Scorzonera) or with corniculate projections at tip — caps [subgenus 
Podospermum (DC.) Lipsch.]. Receptacle pubescent (in some species of 
section Tuberosae) or mostly glabrous, alveolate; alveoles cartilaginous 



'I have deliberately omitted another publication of Bipontinus Schultz "Sceleton 
systematis articulati Cichoriaceanim" (Sch. Bip. in Herb., and Winkl. Jahrb. Pharm. IV, 
1841; vidi tantum seorsum impressum), where a detailed division of Cichoriae is given, but 
without any indication of the rank of the groups used. 



27 

along margins. Ligulate florets yellow, less often red and pink, truncate, 
five-toothed. Anthers sagittate at base, auricles acute or short-setose. 
Style branches slender. Achenes glabrous or pubescent, cylindrical, 
oblong, slightly (very rarely strongly) narrowed at apex; at base 
sometimes with hollow tubular appendage (stalk) (subgenera 
Podospermum (DC.) Lipsch. and Pseudopodospermum (Lipsch. and 
Krasch.) Lipsch.) or without it (subgenus Scorzonerd), ribbed, smooth 
or scabrous along ribs; pappus hairs plumose or barbed, a few (5-10) 
longer, plumose in lower part, barbed above. Glabrous or pubescent 
perennials, less often semishrubs and biennials, with alternate, entire, 
pinnatifid, or pinnatisect leaves. Roots variable in shape, tuberous in 
many species. 

Type of genus: Scorzonera humilis L. 

The genus Scorzonera comprises nearly 170 species, distributed from 
central Europe and the Mediterranean Region to eastern Asia. Members 
of the genus are particularly abundant in Iran, Asia Minor, the Caucasus, 
and Soviet Central Asia, i.e., mostly in arid regions. 

1. Achenes with hollow swollen stalk at base 2. 

+ Achenes at base without hollow swollen stalk. All leaves simple, 

entire, pinnatifid only in S. bicolor. Involucral bracts not 
28 comiculate at tip (subgenus Scorzonerd) 14. 

2. Leaves pinnatisect. Involucral bracts often comiculate (subgenus 
Podospermum (DC.) Lipsch.) 3. 

-I- Leaves simple, undivided. All involucral bracts without 
comiculate projections at tip (subgenus Pseudopodspermum 
(Lipsch. and Krasch.) Lipsch.) 55. 

3. Biennials, with well-developed leafy stems 4. 

+ Perennials usually with poorly developed stems 7. 

4. Involucral bracts with white or yellowish tomentum on outer side 
at base 1. S. lachnostegia (Woron.) Lipsch. 

+ Involucral bracts without tomentum on outer side at base 5. 

5. Lateral segments of leaves elliptical-lanceolate or linear- 
lanceolate, terminal segment large and wide, oblong or ovate- 
lanceolate. Stems mostly ascending or prostrate 

3. S. calcitrapifolia Vahl. 

+ LateraL segments of leaves narrow and long, linear or linear- 



'Treatment by S.J. Lipschitz. 

^Frotn the Italian words scorza — root, and nera — black, because of the outer color of 
the root. Even more probable, from the Spanish name of this plant, originating from 
escorzon — a poisonous snake, as the root is used in Spain as a medicine against snake bite. 



28 

lanceolate, terminal segment little different in shape from lateral 
segments. Stems mostly erect 6. 

6. Achene stalk one-fourth or one-fifth as long as achene. Some 
leaves (particularly cauline ones) undivided; lateral segments few. 
Pappus readily detaching on touch 

4. S. songorica (Kar. and Kir.) Lipsch. and Vass. 

+ Achene stalk one-third or almost half as long as achene. All 

leaves pinnatisect. Pappus not readily detaching on touch 

2. S. laciniata L. 

7. Alpine plants. Stems scape-like. Almost all leaves crowded at 
stem base 8. 

+. Plants of lower and middle zones. Stems leafy 10. 

8. Plants glabrous. Leaves with large obovate, obtuse upper segment, 
and two or three pairs of short linear lateral segments 
(Khevsuretia) 12. S. grigoraschvilii (Sosn.) Lipsch. 

+ Plants more or less pubescent. Upper segment of leaves oblong, 
narrowed toward apex, often acute 9. 

9. Plants green or grayish-green, weakly pubescent. Capitula large, 

1.8-2(3) cm long in full bloom 

10. S. meyeri (C. Koch) Lipsch. 

+ Plants gray from dense pubescence. Capitula smaller, up to 1.5 

cm long in full bloom (Talysh) 

11. S. kirpicznikovii Lipsch. 

10. Achenes (11)12-15(16) mm long. Lateral lobes of leaves very 
narrow, about 1 mm wide (Talysh) 

8. S. grossheimii Lipsch. and Vass. 

29 -I- Achenes shorter 11. 

11. Involucral bracts (outer) usually comiculate at apex. Achenes 
more or less sparsely hairy, less often glabrous. Peripheral achenes 
large, broad to winged, 2-3 mm wide. Lateral lobes of leaves 
rather wide, 2-3(5) nmi wide 

5. S. armeniaca (Boiss. and Huet) Boiss. 

+ Combination of characters different 12. 

12. Stems shorter, 10-25(30) cm high, usually whole plant white- 
tomentose. Achenes bottle-shaped, with ribs more or less 
undulately rugose. Achene stalk 3-4 mm long, with distinct ribs 

9. S. cana (C.A.M.) O. Hoffm. 

+ Stems taller, 20-30(40) cm high, weakly pubescent. Achenes not 
bottle-shaped, ribs not undulately rugose 13. 

13. Rhizome thick, many-headed. Stems virgate, 15-25 cm high. 
Capitula many (10-20 on single plant), narrow-cylindrical, 
relatively small (10-15 nun long and 2-5 mm wide in bloom) 

6. S. schischkinii Lipsch. and Vass. 



29 

+ Rhizome not so thick. Stems slender, branched, not virgate. 
Capitula fewer, obconical (6-8 mm wide in bloom) (Mashhadia) 
7. S. idae (Sosn.) Lipsch. 

14. Plants forming pad-like tussock with numerous stems 15. 

+ Plants not forming pad-like tussock with numerous stems.... 17. 

15. Elastic rubber fibers visible to naked eye on breaking underground 
organs. Achenes p bescent. Capitula large, broadly cylindrical 
or obconical 22. S. tau-saghyz Lipsch. and Bosse 

+ Elastic rubber fibers not visible on breaking underground organs. 
Achenes glabrous. Capitula narrow-cylindrical 16. 

16. Lateral branches usually numerous, flexuous, prickly, without 
capitula. upper leaves filiform. Stems 10-20 cm high 

56. S. acanthoclada Franch. 

+ Prickly lateral branches absent. All leaves linear and basal. Stems 
20-30 cm high 57. S. kuhistanica M. Pop. 

17. Basal and cauline leaves varying in degree of cutting of lamina, 
from almost entire or sharply toothed to pinnatifid. Rorets yellow, 
corolla throat black-purple 19. S. bicolor Freyn and Sint. 

-I- All leaves always entire. Florets yellow, red, pink, single-colored 
18. 

18. Plants with large capitula (especially in fruiting specimens) 
30 reaching (3.5)5-8 cm long and 1.5-2(3) cm wide. Achenes (2)4- 

4.5 cm long, narrowed at apex into beak (section Piptopogon 

C.A. Mey 19. 

-f Plants with smaller capitula (on average, not more than 5 cm 
long in fruiting specimens). Achenes smaller, apex not narrowed 
into beak 25. 

19. Plants small, stems 2-2.5 cm high. Many stems (up to 10) arising 
from root collar. Capitula at fruiting longer than stem 

26. S. albertoregelia Winkl. 

+ Plants much taller, with solitary (or less often two or three) stems. 
Capitula never longer than stem 20. 

20. Capitula surrounded by more or less large adpressed bracts. 
Leaves broadly lanceolate or ovate-lanceolate, (2)2.5-4 cm wide, 
long-acuminate 21. 

+ Capitula without large adpressed bracts. Leaves linear, herbaceous 
or broadly linear (up to 2 cm wide) 22. 

21. Ligulate florets yellow. Bracts usually large, numerous 
(Tadzhikistan) 24. S. bracteosa Winkl. 

+ Ligulate florets pink. Bracts smaller, fewer (Alai) 

25. S. alaica Lipsch. 

22. Florets yellow. Capitula more or less numerous, often in 
corymbose inflorescence (Far East) 23. S. albicaulis Bge. 



30 

+ Florets pink when dried (Soviet Central Asia) 23. 

23. Capitula solitary, terminal on stem or lateral branches. Compound 
inflorescence not forming candelabrum-like corymb. Peduncles 
not always arcuately bent, sparsely leafy 24. 

+ Capitula in more or less compressed candelabrum-like corymb. 

Peduncles arcuately bent, densely leafy 

28. S. petrovii Lipsch. 

24. Plants usually large, up to 30-35 cm high. Involucral bracts not 
black and without black spot 

27. S. tragopogonoides Rgl. and Schmalh. 

+ Plants smaller, 7-15 cm high. Involucral bracts black or with 

distinct black spot 29. S. baldshuanica Lipsch. 

25(18) Root collar fibrous from dead petioles and sheaths of basal leaves 

26. 

+ Root collar not fibrous and usually covered with more or less 

hardened undivided or partly cut up sheaths of basal leaves 

34. 

26. Florets purple or violet-pink. Stems usually densely leafy (section 

Purpureae Lipsch.) 27. 

+ Florets yellow (section Fibrillosae Nakai) 28. 

31 27. Leaves grooved. Stems solitary, with one or two to five capitula 

at apex. Achene ribs smooth. Plants with strong vanilla fragrance 

at time of flowering 20. S. purpurea L. 

-I- Leaves flat. Stems always with single capitulum. Achene ribs in 

upper part toothed-scabrous 20. S. rosea Waldst. and Kit. 

28. Plants of alpine zone, almost leafless or with scape-like lower 
more or less pubescent stems. Capitula large (up to 5 cm long), 
broadly cylindrical. Involucral bracts more or less pubescent (Tien 
Shan) 30. S. subacaulis Lipsch. 

-I- Plants of steppe, forest-steppe and forest zone. Stems well 
developed (sometimes short), glabrous. Involucral bracts glabrous 
29. 

29. Plants small, usually 3-7(10) cm high, with filiform, spiral, narrow 
(1-1.5 mm wide) glaucous leaves, usually longer than stem 
(Trans-Baikal Region) 36. S. curvata (Popl.) Lipsch. 

+ Plants medium-sized (usually more than 10 cm high), with wider 
leaves (more than 1-1.5 mm wide), not longer than stem.... 30. 

30. Leaves ovate-lanceolate or broadly lanceolate, more or less 
distinctly undulate-crimped along margin. Stems mostly branched 
(Crimea and hilly Kazakhstan) 31 S. crispa MB. 

+ Leaves linear, lanceolate-linear or broadly lanceolate, flat along 
margin 31. 

31. Basal leaves with long petioles, broadly lanceolate, up to 4.5-5 



31 

cm wide. Stems mostly tall, up to 35-40 cm high, thick, hollow, 
strongly sulcate, Capitula large, up to 4.5-5 cm long. Plants 

usually with rosettes of numerous basal leaves 32. 

+ Basal leaves with medium-sized petioles, linear-lanceolate, linear, 
or lanceolate, usually narrower than 4.5-5 cm. Stems shorter and 
less thick, indistinctly sulcate. Capitula smaller. Rosettes of basal 
leaves indistinct 33. 

32. Plants green; stems tall, simple (north of the European part of 
the USSR, Urals, eastern Siberia) 

32. S. ruprechtiana Lipsch. and Krasch. 

+ Plants glaucous; stems usually branched (.saline meadows and 
limans of southwestern European part of the USSR and 
Kazakhstan) 32. S. pratorum (Krasch.*) Stankt)v 

33. Capitula narrow-cylindrical. Stem branched, flowering branches 
slender, filiform (Arkadian Mountains in eastern Kazakhstan) .. 

35. S. dianthoides (Lipsch. and Krasch.) Lipsch. 

+ Capitula broadly cylindrical. Stems simple or branched, flowering 

branches thicker 34. S. austriaca Willd. 

34(25) Involucral bracts with red spot at apex 35. 

+ Involucral bracts without red spot at apex 36. 

32 35. Basal leaves linear or linear-lanceolate, scarcely acuminate. 

Capitula (2.5)3-3.5(4) cm long (Siberia) 

42. S. radiata Fisch. 

+ Basal leaves ovate or oval, cuspidate. Capitula (3.5) 4-5 cm long 

(European part of the USSR) 43. S. humilis L. 

36. Plants of saline habitats. Ligulate florets slightly exceeding 
involucre. Pappus snow-white 37. 

+ Plants of other habitats 39. 

37. Root thickened into ovoid-spherical tuber. Young leaves apically 
spirally twisted 59. S. pusilla Pall. 

+ Root not thickened into tuber. Young leaves apically not twisted 
38. 

38. Plants glaucous. Leaves thicker and fleshy. Root collar thickened 
from more or less numerous remnants of hardened sheaths of 
basal leaves; inner surface of sheath lanate. Stems more or less 
numerous, branched. Capitula usually a few (four or five) (eastern 
Kazakhstan) 45. S. mongolica Maxim. 

-I- Plants green. Leaves less thick, somewhat fleshy. Root collar 
not thickened. Stems solitary, usually simple. Capitula solitary, 
or less often two 44. S. parviflora Jacq. 

39. Achenes glabrous 40. 

+ Achenes lanate or villous over entire surface 68. 

40. Plants forming more or less dense tussock. Stems scape-like, 



32 

almost leafless, slender, sometimes erect. Leaves narrowly linear 
or lanceolate, crowded at stem base [section Pulvinares (Boiss.) 

Lipsch.] 41. 

+ Combination of characters different 44. 

41. Stems distinctly leafy. Leaves lanceolate, 4-7 mm wide; basal 
leaves narrowed into grooved, tomentose petiole 

41. S. koslovskyi Sosn. 

+ Stems slightly leafy. Leaves linear, linear-filiform, usually much 
narrower (up to 4 mm wide), not narrowed into grooved petiole 
42. 

42. Plants usually (20)30-40 cm high. Stems mostly branched, 
numerous. Base of aerial part quite densely covered with hardened 
leaf sheaths (Caucasus) 40. S. rigida Auch. ex DC. 

+ Plants shorter, (5)7-15 cm high. Stems simple, few. Base of aerial 
part not covered with hardened leaf sheaths (Caucasus) 43. 

43. Whole plant, particularly leaves and involucres, covered with 
scattered long, flexuous, divergent hairs. Basal leaves linear, 1.5- 

33 2.5 mm wide 39. S. seidlitzii Boiss. 

+ Plants not covered with scattered long, flexuous and divergent 

hairs. Basal leaves flliform-hnear, 0.5-1 mm wide 

37 S. filifolia Boiss. 

44(40) Leaves broadly oblong-ovate, ovate-elliptical, or broadly 

lanceolate, 2-4(8) cm wide, many-veined. Venation arcuate 

45. 

+ Leaves narrower; veins less prominent 46. 

45. Whole plant very densely velutinous (shining), with snow-white 
hairs. Pappus reddish-brown (Akhalkalaki Plateau) 

67. S. ketzkhovelii Sosn. 

+ Whole plant weakly pubescent (not velutinous) or almost glabrous. 

Pappus dirty white (Karatau Range) 

46. S. franchetii Lipsch. 

46. Pappus hairs rigid, blackish-violet 47. 

+ Pappus hairs not blackish-violet 48. 

47. Basal leaves broadly lanceolate or ovate-lanceolate, petiolate 
(Pamiro-Alai) 65. S. hissarica Winkl. 

+ Basal leaves herbaceous, linear, sessile (Turkmenia) 

18. S. raddeana Winkl. 

48. Root thickened into variously shaped tuber 49. 

+ Root not thickened into tuber 59. 

49. Achenes without basal stalk 50. 

+ Achenes with basal stalk 55. 

50. Capitula narrowly cyUndrical, usually numerous (up to 20 and 
more on a plant). Young leaves narrowly linear, apically spirally 



33 

twisted. Achenes smooth or somewhat scabrous 

'. 59. S. pusilla Pall. 

+ Capitula broadly cylindrical, fewer. Basal leaves petiolate, ovate- 
orbicular, or oblong-lanceolate. Achenes strongly scabrous from 
numerous tubercles on ribs. Pappus of snow-white, densely 
plumose hairs 51. 

51. Capitula 4-6 mm wide 52. 

+ Capitula 7-15 mm wide 53. 

52. Stems 20-30 cm high, with long intemodes, leafy up to apex. 
Leaves flat along margin 

60. S. tadshikorum Krasch. and Lipsch. 

-1- Stems 10-15 cm high, scape-like, branched from middle, 
flowering shoots more or less numerous; leaves crowded at base 
of branches; intemodes short, leaves more or less undulate- 
crimped along margin 61. S. ferganica Krasch. 

53. Flowers yellow (Soviet Central Asia) 54. 

34 + Flower pink or violet (Caucasus) 64. S. papposa DC. 

54. Leaves hard, cartilaginous, acuminate, along margin more or less 
undulate-crimped (Turkmenia) 62. S. ovata Trautv. 

-I- Leaves fleshy, obtuse, along margin flat 

63. S. crassifolia Krasch. and Lipsch. 

55(2) Stems more or less tall, sulcate, leafy up to apex. Mature achenes 

straw-yellow. Pappus white 13. S. mollis M.B. 

+ Stems scape-like, usually short, leafy in lower part, less often 

with solitary leaves along whole stem. Pappus dirty gray .... 56. 

56. Leaves bright green (if glabrous), broadly lanceolate, (4)6-12(16) 
mm wide. Tuber fusiform (Kuban) 

16. S. turkeviczii Krasch. and Lipsch. 

+ Leaves glaucous or grayish, narrower, 2-4(6) mm wide. Tubers 
nutty, spherical or ovoid 57. 

57. Flowers red, violet when dry 15. S. suberosa C. Koch 

+ Flowers yellow 58. 

58. Tubers usually borne at soil surface, nutty. Leaves wider, 2-3 
mm wide, often strongly undulate-crimped along margin 

14. S. leptophylla (DC.) Krasch. and Lipsch. 

+ Tubers borne at some depth from soil surface, ovoid, less often 
spherical. Leaves narrower, 1-2 mm wide, usually flat along 

margin 17. S. turcomanica Krasch. and Lipsch. 

59(48) Semishrubs. Base of root collar covered with hardened, entire or 
slightly divided, sheaths of previous year's leaves. Stems 

numerous, virgate (Tien Shan) 

58. S. pseudodivaricata Lipsch. 

-I- Perennials. Stems numerous, not virgate 60. 



34 

60. Flowers orange (mountains of Soviet Central Asia) 

49. S. transiliensia M. Pop. 

+ Flowers yellow or pink 61. 

61. Leaves narrow, linear, 1-7(10) mm wide 62. 

+ Leaves broadly lanceolate or ovate-lanceolate, wider, up to 2- 

2.5 cm wide 66. 

62. Whole plant white tomentose. Ligulate florets pink (Shugnan). 

50. S. gracilis Lipsch. 

+ Plants grayish pubescent or glabrous. Ligulate florets yellow ... 
63. 

63. Achenes smooth along ribs. Capitula in racemes, usually few- 
flowered 64. 

35 + Peripheral achenes with acute or obtuse tubercles along ribs in 
upper part. Capitula many-flowered 65. 

64. Capitula in long, sometimes interrupted racemes, often on short 
peduncles, sometimes almost sessile, narrowly cylindrical, bearing 
6-8(12) flowers. Raceme more or less compact. Capitula before 
flowering ellipsoid or cylindrical. Involucral bracts 12-15 (Soviet 
Central Asia) 47. S. turkestanica Franch. 

+ Capitula in indistinct corymbose raceme, usually on longer 
peduncles, broadly cylindrical. Raceme often lax. Capitula before 
flowering globose. Involucral bracts 20-32 (Soviet Central Asia) 
48. S. iliensis Krasch. 

65. Stems 20-50 cm high, solitary, straight, branched in upper half 
with projecting, erect, slender peduncles. Leaves flat along 
margins 51. S. stricta Homem. 

+ Stems shorter, 8-20 cm high, mostly two or three, usually 
branched from base, with bent (less often erect), thicker peduncles. 
Leaves somewhat hairy along margins 52. S. pubescens DC. 

66. Plants taller, 25-75 cm high. Stems usually branched in upper 
part or less often from middle. Leaves mostly crowded in lower 
third of stem, mostly flat along margins. Capitula broadly 
cylindrical 53. S. hispanica L. 

-I- Plants shorter, (5)10-25(30) cm high. Stems usually branching 
from base. Leaves borne along entire stem, often undulate-crimped 
along margin 67. 

67. Leaves broadly lanceolate, strongly attenuate, acuminate. Achenes 
along ribs tuberculate-scabrous 

54. S. inconspicua Lipsch. ex Pavl. 

-f- Leaves obovate-lanceolate, slightly acuminate. Achenes along 

surface squamose-prickly 55. S. chantavica Pavl. 

68(39) Roots cylindrical or fusiform, not thickened into ovoid-spherical 
tuber. Plants usually large 69. 



35 

+ Roots thickened into ovoid-spherical tuber. Usually small desert 
plants of spring (ephemerals) 76. 

69. Plants snow-white from dense velutinous pubescence. Leaves 
elliptical or broadly lanceolate, with many pubescent veins 
(Akhalkalaki) 66. S. sosnovskyi Lipsch. 

-I- Nature of pubescence different 70. 

70. Leaves wide, (1.5)2-5 cm wide, broadly lanceolate or lanceolate 

71. 

+ Leaves narrow, linear, 1-5 mm wide 72. 

7 1 . Gray, less often whitish plants with appressed hairs. Inflorescence 
36 paniculate. Leaves along entire margin distinctly cartilaginous, 

scabrous 68. S. latifolia (Fisch. and Mey.) DC. 

+ Whole plant densely tomentose (but not velutinous). Inflorescence 
almost corymbose (karabakh) 69. S. pulchra Lorn. 

72. Leaves with many veins, linear, 3-5(8) mm wide 

70. S. safievii Grossh. 

+ Leaves with one to three veins 73. 

73. Plants with scape-like solitary stems, (5)10-15 cm high. Leaves 
filiform, up to 1 mm wide. Achenes pubescent, but not lanate 

38. S. lipskyi Lipsch. 

+ Combination of characters different 74. 

74. Stems virgate, upright, straight. Basal leaves soft, without distinct 
spiny cusp. Capitula few-flowered (with six or seven florets). 
Achenes 9-10 mm long (excluding pappus) 

73. S. ramosissima DC. 

-I- Stems ascending or erect, not virgate. Leaves harder, mostly with 
prominent veins, long-attenuate into very thin, often bent tip. 
Capitula many-flowered (with more than 20 florets). Achenes 5- 
7 mm long (excluding pappus) 75. 

75. Involucre puberulent or almost glabrous. Outer involucral bracts 
without subulate awn, less than half as long as inner bracts 
(Caucasus) 72. S. biebersteinii Lipsch. 

+ Involucre densely lanate-villous. Outer involucral bracts at least 
half as long as inner ones (usually longer than half), with distinct 
subulate awn (south of the European part of the USSR, western 

Siberia, Soviet Central Asia) 71. S. ensifolia M.B. 

76(68) Stems branched (Caspian deserts, Soviet Central Asia, western 
Siberia) (section Tuberosae Lipsch.) 77. 

-I- Stems simple (Caucasus) (subsection Lanatae Lipsch.) 83. 

77. Capitula drooping after blooming. Leaves hard, almost coriaceous, 
with prominent veins 78. 

+ Capitula somewhat drooping after flowering, leaves soft, not 
coriaceous. Veins less prominent 81. 



36 

78. Leaves broadly lanceolate, 0.8-3 cm wide, with many (5-11), 
mostly rather prominent veins 79. 

+ Leaves narrowly lanceolate or linear, 0.3-0.5 cm wide, with three 
to five less prominent veins 80. 

79. Leaf margins flat 79. S. gageoides Boiss. 

+ Leaf margin somewhat strongly undulate-crimped (sometimes 

rolled); their tips often spirally twisted 

80. S. circumflexa Krasch. and Lipsch. 

80. Involucre 8-10 mm long. Aerial part [of plant] 5-7 cm high. 
Capitula often solitary. Tuber very irregular, with large tubercles 
especially in lower part 82. S. oxiana M. Pop. 

+ Involucre 10-15 mm long. Aerial part [of plant] 10-20(30) cm 
37 high. Capitula numerous. Branches more or less robust, ascending, 

often virgate after fruiting 

81. S. litvinovii Krasch. and Lipsch. 

81. Leaves linear, 2-5 mm wide 82. 

+ Leaves broadly lanceolate or oblong-lanceolate, 0.5-0.9(2.1) cm 

wide, covered with appressed tomentose hairs 

77. S. sericeo-lanata (Bge.) Krasch. and Lipsch. 

82. Plants small, 4-8.5(11) cm high. Leaves canescent or almost 
glabrous 76. S. tuberosa Pall. 

-I- Plants taller, 8.5-20(40) cm high. Leaves and stems covered with 

more or less divergent silky hairs 

77. S. sericeo-lanata (Bge.) Krasch. and Lipsch. 

83(76) Plants taller, (10)15-20(25) cm high with ascending stems, leafy; 
leaves usually much shorter than stems. Capitula large, up to 2- 
2.5 cm long. Florets one and one-half to two times as long as 

involucre, lemon-yellow 

74. S. lanata (L.) Hoffm. 

-I- Plants less tall, 3-7(10) cm high, with drooping (during fruiting) 
stems; leaves almost as long as stems. Capitula smaller, 0.8-1 
cm long. Florets scarcely longer than involucre, orange-yellow 
75. S. pseudolanata Grossh. 

Subgenus I. Podospermum (DC.) Lipsch. Fragm. Monogr. 
Scorzonera I (1935) 7.— Gen. Podospermum DC. Fl. Fr. ed. 3, IV (1805) 
61; Cass, in Diet. sc. nat. XLII (1826) 77; Less. Synops. Compos. (1832) 
133 (sub nom. Podosperma, non Labill. excl. syn. Roth and Moench); 
DC. Prodr. VII (1838) 110; Endlicher, Gen. pi. (1838) 497.— Subgen. 
Eupodospermum Rouy, Fl. Fr. X (1908) 16; Ej. Conspect. Fl. Fr. (1927) 
170. — Sect. Podospermum Benth. in Benth. and Hook. f. Gen. pi. II 
(1873) 532; Boiss. H. or. Ill, 757; O. Hoffm. in Pflanzenfam. IV, 5 
(1893) 365. — Achenes at base with hollow swollen stalk. Leaves 



37 

pinnatisect. Involucral bracts often with comiculate projections (caps) 
at tip. 

Type of subgenus: S. laciniata L. 

Series 1. Laciniatae Lipsch. in Addenda XXVIII, 717. — Biennials 
with well-developed leafy stems. Achenes cylindrical, glabrous. 

1. S. lachnostegia (Woron). Lipsch. Fragm. Monogr. Scorzonera II 
(1939) 11. — Podospermum lachnostegium Woron. in Tr. Bot. Inst. Akad. 
Nauk SSSR, Ser. 1, 1 (1933) 223; Grossh. Fl. Kavk. IV (1934) 234; 
Grossh. Opred. Rast. Kavk. 507. 

Biennial? Stems up to 25 cm high, more or less pubescent, branched 
from base. Leaves sessile, pinnatisect; lateral leaf segments linear or 
linear-lanceolate, with elongate terminal segment. Lateral peduncles 
longer than leaves, leafless or with one or two linear, acuminate, small 
38 leaves. Involucral bracts on outer side densely white lanate; inner bracts 
oblong-lanceolate, two times as long as outer bracts, almost glabrous; 
all bracts with distinct midrib. Achenes 9-10 mm long, cylindrical, 
glabrous; pappus snow-white, as long as achene, of plumose bristles. 
Flowering May. 

Caucasus: Ciscaucasia. Endemic. Described from the vicinity of 
Zapadinsky farmstead. Type in Leningrad. 

Note. It differs from all species of subgenus Podospermum by having 
floccose outer involucral bracts. 

2. S. laciniata L. Sp. pi. (1753) 791; Willd. Sp. pi. Ill, 3 (1803) 
1506; Boiss. Fl. or. Ill, 757 (excl. p. calcitrapifolia); Schmalh. Fl. II, 
141, p.p.; Krasch. in Fl. YugOrVost. VI, 448; Askerova in fl. Azerb. 
VIII, 515 p. max. p.; Lipsch. in Fl. Turkm. VII, 302; Papava in Fl. 
Gruzii, VIII, 637. — S. laciniata var. subintegrifolia Lipsch. in Spisok 
Rast. Gerb. Fl. SSSR XIII (1955) 131.— 5. octangularis Willd. op. cit. 
Ill, 3 (1803) 1506.— Podospermum laciniatum (L.) DC. R. Fr. IV (1805) 
62; DC. Prodr. VII, 1, HI, (excl. var. subulatum); Ldb. R. Ross. II, 2, 
782; Grossh. Fl. Kavk. IV (1934) 234; Grossh. Opred; Rast. Kavk. 507; 
Kryl. Fl. Zap. Sib. XI, 2979, p. min. p. — P. muricatum DC. Syn. pi. in 
Fl. Gall. (1806) 265.— Exs.: GRF No. 3997. 

Biennial. Roots usually slender. Plant more or less arachnoid 
pubescent, especially in lower part, later often almost glabrescent. Stems 
erect or less often ascending, bent, (10)20-40(50) cm high, solitary or 
several, sulcate, simple or branched. Basal leaves long-petiolate, broader 
at base, pinnatisect, with narrowly linear or linear-lanceolate lateral 
segments; terminal segment longer than lateral ones; linear-lanceolate 



38 

or lanceolate; cauline leaves reduced, short-petiolate; number of lateral 
segments highly variable. Capitula cylindrical, 6-8 mm wide at flowering, 
10-15(20) mm wide at fruiting. Involucre sparsely arachnoid pubescent, 
glabrescent; outer involucral bracts deltoid-lanceolate, sometimes 
comiculate at apex, with more or less distinct midrib; inner bracts oblong- 
lanceolate, two to three times as long as outer ones; all bracts narrow- 
scarious along margin. Florets pale yellow, longer than involucre. 
Achenes 10-12 mm long, cylindrical, light gray, glabrous, smooth, with 
obtuse longitudinal ribs, slightly bent in lower part (approximately one- 
third to half the length), swollen toward base, and light colored. Stalk 
straw-yellow, 4 mm long. Flowering April to May. 

Plains and foothills on clayey and stony soils, sometimes as weed. — 
European Part: Crimea, Black Sea Region, Lower Don, Lower Volga; 
Caucasus: Ciscaucasia, Eastern and Southern Transcaucasia, Dagestan; 
Soviet Central Asia: mountainous Turkmenia. General distribution: 
central and southern Europe, Balkan- Asia Minor, Armenia and Kurdistan, 
Iran. Described from Germany and France. Type in London. 

Note. A highly polymorphic species, which should undoubtedly be 
divided into several local races. Insufficient material, however, does not 
permit us to do this at present. Within the species, var. elongata Sch. 
39 Bip. is distinguished by erect, tall (to 60 cm), thickened (to 7-8 mm), 
knotty, hollow, scabrous stem, usually with scattered fine papillae, very 
long (to 20-25 cm) leaves, and longer ray florets. Evidently, this variety 
is found under conditions of high humidity; in the USSR, such plants 
are known from Armenia (for example, the collection of J.N. Woronow 
from the vicinity of Takyaltu, in crop field. No. 12680). Probably, the 
name S. octangularis Willd. applies particularly to this variety. 

3. S. calcitrapifolia Vahl, Symb. 2 (1791) 87; Willd. Sp. pi. Ill, 3 
(1803) 1505; Guss. PI. rar., 323. — Podospermum calcitrapifolium (Vahl) 
DC. Fl. Fr. vol. VI (1815) 455; C.A.M. Verzeichn. (1831) 62, p. p.; 
DC. Prodr. VII, 1, 110; Grossh. Fl. Kavk. IV (1934) 234; Papava in Fl. 
Gruzii, VIII, 638.— Z'. intermedium Guss. in DC. Prodr. VII, 1 (1838) 
110, p. p.— P. buxbaumii C. Koch in Linnaea XVII (1843) 214.— P. 
heterophyllum C. Koch Ibid (1843) 275. — S. laciniata p. calcitrapifolia 
(Vahl) Bisch. ex Boiss. Fl. or. Ill (1875) 757. 

Biennial. Plant glabrous or more or less arachnoid pubescent. Stems 
several or solitary, (15)20-30(45) cm high, branched above, usually 
ascending or prostrate below. Leaves pinnatisect or pinnatifid with linear- 
lanceolate, oblong-elliptical, or obovate-lateral segments and larger 
terminal segment; basal leaves long-petiolate, broader at base; cauline 
leaves reduced toword stem apex, short-petiolate or sessile; leaf incisions 
and shape of lateral segments variable. Capitula solitary, terminal on 



39 

stems and branches, cylindrical, 5-8 mm wide at flowering, 1-1.5(2) 
cm at fruiting. Involucre initially whitish pubescent, glabresent; involucral 
bracts with narrow scarious margin and distinct midrib; outer bracts 
smaller, deltoid-lanceolate; inner ones oblong-lanceolate, two to three 
times as long as outer bracts; ligulate florets yellow, slightly longer 
than involucre. Achenes 10-12 nrni long, grayish, glabrous, cylindrical, 
lower part twisted and broadened into stalk, with longitudianal light- 
colored ribs; ribs glabrous in stalk region, and indistinctly pubescent or 
(in immature achenes) glabrous above; stalk light colored, up to 4 mm 
long, one-third to half as long as achene; pappus as long as achene, 
white or reddish at base, pappus bristles plumose, longer hairs scabrous 
toward apex. Flowering April to May. 

Plains and foothills, on clayey and stony soils, sometimes in crop 
fields as weed. — Caucasus: Ciscaucasia,? Dagestan, Eastern and Southern 
Transcaucasia. General distribution: Mediterranean Region, Iran. 
Described from Tunisia. Type in Copenhagen (?). 

Note. Pending the collection of more complete and abundant 
material, it is not clear whether Scorzonera calcitrapifolia deserves to 
be recognized as a separate species. Possibly, it should be treated as a 
40 variety of S. laciniata; that is how this question was interpreted by 
Boissier (op. cit.). The only dependable distinction between the two 
species is the presence of a wider and larger terminal leaf segment in S. 
calcitrapifolia. However, this character is variable, and in several cases 
transitional forms are found that are difficult to include in either of the 
two species in question. The geographic distribution of S. calcitrapifolia 
also is not clear, as the study material of this species at my disposal 
was extremely limited. To judge from the analysis of type material of 
Podospermum buxbaumii C. Koch, which I found in the Herbarium of 
the Botanical Institute, Academy of Sciences of the USSR, the latter 
species is absolutely identical with Scorzonera calcitrapifolia Vahl, to 
the synonyms of which it has been relegated. 

4. S. songorica (Kar. and Kir.) Lipsch. and Vass. in Spisok Rast. 
Gerb. Fl. SSSR, XIII (1955) 131; in Fl. Turkm. VII (1960) 302; 
Kovalevskaja in Fl. Uzb. VI, 436. — Podospermum laciniatum var. 
songoricum Kar. and Kir. in Bull. Soc. Nat. Mosc. XV, 2 (1842) 396. — 
P. laciniatum var. integrifolium Ldb. Fl. Ross, II, 2, (1845-1846) 782. — 
P. laciniatum auct. fl. Asiae Mediae, p. max. p. — P. songoricum (Kar. 
and Kir.) Lipsch. and Vass. in sched. — S. praticola Bomm. in sched. herb. 
"Bommiiller: Iter Persico-turcicum 1892-1893, No. 4109" nom. nud. 

Biennial. Plant green; almost glabrous or weakly arachnoid- 
pubescent. Stems few, usually branched, less often simple, (10)15^0(50) 
cm high. Leaves pinnatisect, sometimes (especially often cauline) 



40 

undivided, oblong-linear or linear; lateral segments usually a few, linear, 
linear-lanceolate or oblong; terminal segment longer, oblong or oblong- 
linear, gradually narrowed below; basal leaves petiolate; cauline leaves 
sessile, narrowed toward base. Capitula numerous, 5-6 mm wide at 
flowering, up to 1 cm wide at fruiting. Involucre arachnoid-pubescent 
initially, almost glabrous at fruiting; outer involucral bracts oblong-ovate, 
sometimes corniculate at apex, one-third to half as long as oblong 
lanceolate inner bracts; ligulate florets yellow. Achenes narrowly 
cylindrical, almost straight, on lower side weakly thickened below, 
grayish, 10-12 mm long, with smooth longitudinal ribs; stalk light 
colored, 2-2.5(3) mm long, one-fifth to one-fourth as long as achene; 
pappus readily detaching, dirty white, as long as or slightly shorter than 
achene, bristles plumose. Flowering June to July (Plate II, Fig. 2.) 

Montane steppes, steppe meadows in river valleys, as weed in 
irrigated and nonirrigated fields, sometimes on slightly saline habitats. — 
Soviet Central Asia: mountainous Turkmenia, Tien Shan, Pamiro-Alai, 
Dzhungaria-Tarbagatai, Balkhash Region, Aralo-Caspian. General 
distribution: Dzhungaria-Kashgaria (Kuldzha). Described from the Lepsa 
River. Type in Leningrad. 

Series 2. Armeniacae Lipsch. in Addenda XXVIII, 717.— Perennials. 
Achenes often hairy, dimorphic: outer achenes broadened and winged, 
inner ones narrower and wingless. 

41 5. S. armeniaca (Boiss. and Huet) Boiss. Fl. or. Ill (1875) 760.— 

Podospermum armeniacum Boiss. and Huet in Boiss. Diagn. pi. or. ser. 
II, No. 3 (1856) 92.— P. isthmocarpum Vass in Grossh. Opred. Rast. 
Kavk. (1949) 508. — P. pallasianum (Krasch. and Lipsch.) Grossh. Ibid 
(1949) 5Q%.—Scorzonera pallasiana Krasch. and Lipsch. in Fl. Azerb. 
VIII (1961) 516 (descr. ross.). 

Perennial. Plant greenish, pubescent,' especially in lower part, 
glabrescent. Stems simple or branched, 15-30(40) cm high, usually two 
to two and one-half times as long as basal leaves. Leaves pinnatisect 
with linear or linear-lanceolate lateral segments, with undulate margin, 
and longer linear, linear-lanceolate or oblong terminal segment; basal 
and lower cauline leaves petiolate, petioles broadened toward base; upper 
cauline leaves reduced, often entire. Capitula solitary, terminal on stem 
and branches, large, 1-1.5(2) cm long and 6-8 mm wide. Involucre 
arachnoid-pubescent or almost glabrous; outer bracts usually corniculate 
at apex, oblong-ovate or lanceolate, much shorter than oblong-lanceolate 
inner bracts; margin, especially with fruits, scarious; ligulate florets 
yellow, bluish on outer side when dry. Achenes 7-10 mm long, with 
longitudinal ribs, more or less scattered pubescent above, usually 
dimorphic: outer achenes oblong, up to 3.5 mm wide, with more or less 



41 

broad-winged lateral ribs (other ribs less prominent), more or less 
scarious-rugose; inner achenes cylindrical, narrower, wingless; achene 
stalk 2-3 mm long; pappus slightly longer than achenes, white, pappus 
bristles plumose. Flowering April to June. 

Dry montane, stony, and clayey slopes, sometimes as weed. — 
Caucasus: Eastern and Southern Transcaucasia, Talysh. General 
distribution: Iran. 

5. armeniaca (Boiss. and Huet) Boiss. was described from eastern 
Anatolia. Type in Geneva. 

Note. The type of S. armeniaca is not known to me. My description 
of the species is based on Caucasian specimens and differs slightly from 
that of Boissier. However, I have no doubt about the similarity of S. 
armeniaca with S. pallasiana. The characteristic feature of both taxa is 
the pubescent achene — a character very rarely found in the members of 
subgenus Podospermum (DC.) Lipsch. S. pallasiana Krasch. and Lipsch. 
was described in Russian from the Nakhichevan ASSR; the type is in 
Leningrad. Podospermum isthmocarpum Vass. (in diagnosis distinguished 
by "unique bottle-shaped achenes, swollen in the middle and with a 
constriction"), with its pubescent seeds, possibly belongs to S. armeniaca 
(Boiss. and Huet) Boiss. The bottle-shaped swelling of the achenes in 
this species is explained by the presence of a parasite in them, which 
became apparent from my sections. 

Series 3. Schischkinianae Lipsch. in Addenda XXVIII, 717. — 
42 Perennials. Root collars knotty. Stem well developed, leafy. Capitula 
small (up to 8 mm wide at flowering). Achenes glabrous, cylindrical. 
Growing in middle and lower mountain zones. 

6. S. schischkinii Lipsch. and Vass. sp. nova in Addenda XXVIII, 
717. — Podospermum schischkinii Vass. in Grossh. Opred. Rast. Kavk. 
(1949) 508 (descr. ross.). 

Perennial. Rhizome thick, many-headed; whole plant almost glabrous 
or weakly arachnoid-pubescent, mainly in lower part. Stems (10)15-25 
cm high, numerous, slender, virgate, branched, leafy. Leaves pinnatisect; 
basal and lower cauline leaves numerous, with petioles broadened into 
sheaths, remnants of which densely cover root collar; lateral segments 
of leaves few, usually long, narrow (1 mm wide); upper cauline leaves 
reduced, sessile, undivided, slightly broadened and semiamplexicaul at 
base. Capitula numerous, solitary, terminal on stem and branches, 
parrowly cylindrical, 10-15 mm long and 'i-A mm wide (at flowering). 
Involucre arachnoid-pubescent; outer involucral bracts oblong-ovate; inner 
ones oblong-lanceolate, two to two and one-half times as long as outer 
ones. Ligulate florets yellow, slightly longer than involucre, ligules 
fimbriate at apex (over 0.5-0.7 mm length). Achenes 9-10 mm long. 



42 




Plate II. 



1 — Scorzonera meyeri (C. Koch) Lipsch.; 2 — S. songorica (Kar. and Kir.) 
Lipsch. and Vass. 



43 

glabrous, narrowly cylindrical, with longitudinal ribs, base clavately 
thickened and slightly curved, stalks 1.5-1.75 mm long; pappus dirty- 
white, as long as achene, pappus bristles plumose. Flowering June to 
July. 

Stony slopes near seacoast. — Caucasus: Western Transcaucasia 
(Anapa). Endemic. Described from Anapa. Type in Leningrad. 

7. S. idae (Sosn.) Lipsch. comb. nova. — Podospermum idae Sosn. 
in Dokl. Akad. Nauk Azerb. SSR, IV, 5 (1948) 211; Grossh. Opred. 
Rast. Kavk. 508; Papava in Fl. Gruzii VIII, 641.— Ic: Fl. Gruzii VIII, 
Plate 433. 

Perennial. Plant glaucescent. Stem up to 25 cm high, simple or 
mostly branched almost from base with apically adpressed branches, 
leafly, weakly pubescent. Leaves pinnatisect; basal leaves up to 15 cm 
long, simple, narrowly linear, sometimes with a few short linear segments 
in upper part; lower and upper cauline leaves similar to basal leaves, 
narrow-linear, reduced. Capitula obconical, small, 8 mm wide at 
flowering, about 15 mm long and 10 mm wide at fruiting. Involucre 
weakly pubescent; outer bracts short, middle bracts lanceolate, inner 
ones oblong-lanceolate; all bracts sUghtly scarious along margin. Ligulate 
florets yellow, one and one-half times as long as involucre. Achenes 8 
mm long, glabrous, outer achenes appressed, slightly broadened, angular- 
44 ribbed; stalk white, 2 mm long; pappus white, almost as long as achene 
or slightly longer, bristles soft. Flowering July. 

Dry slopes in middle mountain zone. — Caucasus: Eastern 
Transcaucasia (Mashhadia). Endemic. Described from Mashhadia. Type 
in Tbilisi. 

Series 4. Grossheimianae Lipsch. in Addenda XXVIII, 718. — 
Perennials with well-developed stems. Leaf sheaths covering root collar. 
Achenes slender, terete, long (up to 16 mm). Growing in middle mountain 
zone. 

8. S. grossheimii Lipsch. and Vass. sp. nova in Addenda XXVIII, 
718. — Podospermum grossheimii Vass. in Grossh. Opred. Rast. Kavk. 
(1949) 509 (descr. ross.). — Scorzonera grossheimii (Vass.) Lipsch. ex 
Isaev in Fl. Azerb. VIII (1961) 519 (descr. ross.). 

Perennial. Plant weakly arachnoid-pubescent, glabrescent, with knotty 
root. Stems numerous, erect or ascending, slender, branched in lower 
part, (15)20-25 cm high. Leaves pinnatisect, predominantly basal or 
crowded in lower part of stem; lateral segments of leaves narrowly linear 
or linear-lanceolate, about 1 mm wide, remote; terminal segment oblong; 
basal and lower cauline leaves with long petioles broadened at base 
into straw-yellow to brown sheath, covering root collar; cauline leaves 
reduced, solitary, undivided, narrowly linear, sessile. Capitula narrowly 



44 

cylindrical, solitary, terminal on stem and branches, 4-5 mm wide at 
flowering, up to 1 cm wide at fruiting. Involucre arachnoid-pubescent, 
glabrescent at fruiting; involucral bracts oblong-lanceolate or lanceolate; 
outer bracts often comiculate at apex; inner bracts two to three times as 
long as outer ones. Ligulate florets yellow. Achenes (11)12-15(16) mm 
long, glabrous, smooth, slender, cylindrical, grayish-reddish-brown, with 
longitudinal ribs; stalks light colored, one-fourth as long as achene; 
pappus dirty white, its bristles plumose, longer ones barbed-scabrous 
above. Flowering June to July. 

Dry rubble slopes. — Caucasus: Talysh. Endemic. Described from 
Talysh. Type in Leningrad. 

Series 5. Canae Lipsch. in Addenda XXVIII, 718. — Perennials. 
Achenes in lower part truncate and broadened, "bottle-shaped"; outer 
achenes wider; inner ones narrow, terete. Growing in foothills and foothill 
plains. 

9. S. cana (C.A.M.) O. Hoffm. in Pflanzenfam. IV, 5 (1893) 365; 
Askerova in Fl. Azerb. VIII, 516. — S. cana (C.A.M.) Grossh. in Beih. 
Bot. Centralbl. XLIV, 2 (1928) 2A1 .—Podospermum canum C.A.M. 
Verzeichn. (1831) 62, p. p. quoad plant, becuens; DC. Prodr. VII, 1, 
110-111, excl. var. glabratum; Ldb. Fl. Ross. II, 2, 781, excl. var. 
glabratum; Grossh. Fl. Kavk. IV (1934) 234, p. p.; Papava in Fl. Gruzii 
45 VIII, 641, p. p. — S. laciniata auct. non L.: M.B. fl. taur-cauc. II (1808) 
237 excl. syn. Jacq.; Ill (1819) 524, p. p. — Scorzonera runcinata Pall. 
Reise in die siidl. Statth. Russ. Reichs. I (1779) 221, nomen; Krasch. in 
Fl. Yugo-Vost. VI (1936) 448 (descr. ross.). — Arachnospermum canum 
(C.A.M.) Domin in Ber. Deutsch. Bot. Gesellsch. LVIIIa (1940) 263.— 
Ic: Fl. Azerb. VIII, Plate LIII, 1. 

Perennial. Plant grayish from arachnoid-pubescence, glabrescent. 
Stems ascending, weak, usually few, (7)8-12(20) cm long, simple or 
branched. Leaves mostly in lower part, pinnatisect with narrowly linear 
or lanceolate (and oblong)-linear segments; these also broadly lanceolate 
or oblanceolate; terminal segment oblong; basal leaves long-petiolate, 
petiole broadened at base into light colored glossy sheath, covering root 
collar; cauline leaves reduced, sessile, sometimes undivided. Capitula 
(1)1.2-1.5(2) cm long, 5-10 mm wide (at flowering), solitary, terminal 
on stems and branches. Involucre arachnoid-pubescent, glabrescent; outer 
involucral bracts lanceolate, often comiculate at apex; inner bracts 
oblong-lanceolate, two to two and one-half times as long as outer ones, 
margin more or less scarious. Ligulate florets yellow, sometimes turning 
pink on drying, longer than involucre. Achenes (7)9-10 mm long, 
longitudinally ribbed, in lower part truncate, and broadened, with 
prominent ribs; outer achenes broader, inner ones narrower, slender, 



45 

cylindrical; stalk almost one-third to half as long as achene; pappus 
dirty white, as long as achene, pappus bristles sparse and long plumose, 
brownish in lower part. Flowering April to June. 

Dry rubbly-stony slopes, foothills and plains. — European Part: 
Crimea, Lower Don, Lower Volga; Western Siberia: ?southem part of 
Upper Tobol; Caucasus: Ciscaucasia, Eastern Transcaucasia, Dagestan. 
General distribution: Iran (northern). Described from the Caucasus 
(vicinity of Baku). Type in Leningrad. 

Note. S. cana is a polymorphic species and, after collection of 
additional new material, probably will be divided into several local races. 
Meyer, under the name used by him, Podospermum canum, mixed up 
the plain and mountain forms. I designate as the type of this species the 
less pubescent specimens collected by Meyer "In collibus pr. Baku 
12. IV. 1830." The specimens from Talysh, also determined by Meyer 
as Podospermum canum, are strongly canescent, belonging to the high- 
mountain members of subgenus Podospermum, representing a unique. 
Talysh race, which I am separating under the name S. kirpicznikovii 
m. among the study material of S. cana is a sharply differentiated variety 
with short, 6-7(8) mm, wider bottle-shaped achenes, the achene stalk 
being brown with prominent white ribs, the ribs in the upper part of 
the fruit hairy-rugose — var. verucoso-costata Lipsch. — Specimens from 
northern Kazakhstan (vicinity of Petropavlovsk, 6,VI,1844, K. 
46 Meinsgauzen) are distinguished by greater height (25-30 cm), erect stems, 
almost entire, linear, basal leaves, some of them pinnatisect, and the 
root collar more densely covered with leaf sheaths. The achenes are 
unknown. It evidently is a separate race requiring further study in nature. 

Series 6. Alpigenae Lipsch. in Addenda XXVIII, 719. — Ser. 
Alpigenae Grossh. generis Podospermi DC. in Zam. po Sist. i Geogr. 
Rast. Tbil. 13 (1947) 66. — Forming tussock. Stem scape-like with a few 
reduced leaves. Leaves predominantly basal, Capitula large, 1.5-2.5 cm 
long (at flowering). Outer involucral bracts usually comiculate. Achenes 
slender, cylindrical, glabrous. Growing in Caucasian high mountains. 

10. S. meyeri (C. Koch) Lipsch. in Fl. Azerb. VIII (1961) 518.— 
Podospermum meyeri C. Koch in Linnaea XXIII (1850) 659; Grossh. in 
Zam. po Sist. i Geogr. Rast. Tbil. 13 (1947) 66; Grossh. Opred. Rast. 
Kavk. 508; Dmitrieva, Opred. Rast. Adzharii, 287; Papava in Fl. Gruzii 
VII, 638.— P. canum p. glabratum DC. Prodr. VII, 1 (1838) 111; Ldb. 
Fl. Ross. II, 2, 163.— P. ponticum Stev. in Bull. Soc. Nat. Mosc. XXIX, 
4 (1856) 404 in textu. — P. canum var. integrifolia Trautv. in Tr. Peterb. 
Bot. Sada II (1873) 558. — Scorzonera jacquiniana (C. Koch) Boiss. p. 
p. and p. alpina Boiss. p. p. in Fl. or. Ill (1875) 757. — Podospermum 
alpigenum auct.: Grossh. Fl. Kavk. IV (1934) 234; Grossh. op. cit. (1947) 



46 

67; Grossh. Opred. Rast. Kavk. 508; Dmitrieva, op. cit. 287; Papava, 
op. cit. 641. — P. tenuisectum Grossh. and Sosn. in Grossh. op. cit. (1947) 
67, excl. plant, talyschens. — IP. alpigenum C. Koch in Linnaea XXIII 
(1850) 658. — IScorzonera alpigena (C. Koch) Groosh. in Beih. Bot. 
Centralbl. XLIV, 2 (1928) 247. 

Perennial. Plant green or greenish-gray, more or less pubescent or 
often glabrous plants with thick root; root collar branched, covered with 
brown sheaths of slightly divided basal leaves; stems scape-like, (3)5- 
15(20) cm high, few, less often solitary. Leaves mostly basal, half as 
long to as long as stem, pinnatisect or entire (var. integrifolia Trautv.), 
narrowed into petiole basally broadened into sheath; lateral segments of 
leaves remote, usually less numerous, linear, acuminate; terminal segment 
oblong, linear; cauline leaves reduced, few, two or three (five), linear, 
often undivided. Capitula solitary at apices of stem and less often of 
branches, large, (1.5)1.8-2(2.5) cm long and 1-1.2 cm wide (at 
flowering). Involucre pubescent, later glabrescent. Involucral bracts often 
dark colored, especially at apex; outer bracts lanceolate, often corniculate 
at apex, one-third to half as long as oblong-lanceolate inner bracts. 
Ligulate florets yellow (in dry condition light lilac), distinctly (one and 
one-half to two times) longer than involucre. Achenes slender, terete, 
9-10 mm long, with longitudinal thin ribs, at base with light colored, 
47 slightly bent, 1(2) mm long stalk; pappus slightly shorter than achene 
or as long, rusty or dirty white, pappus bristles sparsely plumose, finely 
barbed above. Flowering June to August. (Plate II, Fig. 1). 

In alpine zone on meadows, grass plots, screes, moraines, at 2,400- 
2,900 m. — Caucasus: Ciscaucasia (eastern), Dagestan, Eastern and 
Southern Transcaucasia. General distribution: Balkans-Asia Minor, 
Armenia and Kurdistan, Iran. Described from Elbrus. Type in Leningrad. 

Note. Under the name 5. meyeri (C. Koch) Lipsch. I am uniting 
three "species" of Caucasian alpine "Podospermum, " which were 
separated by Grossheim: Podospermum meyeri, P. tenuisectum, and P. 
alpigenum. Specimens of Guriel (in the Herbarium of the Botanical 
Institute, Academy of Sciences of the USSR), identified as Podospermum 
canum C.A.M. var. glabratum DC, and cited by DeCandolle (cp. Prodr. 
VII, 1 (1838) 111), correspond to the interpretation of C. Koch [cp. 
Linnaea, XXIII (1850) 659] of his species Podospermum meyeri. As 
regards the species Podospermum alpigenum C. Koch, S. alpigena (C. 
Koch) Grossh. (cp. Beih. Bot. Centralbl. XLIV, 2 (1928) 247, where for 
the first time this combination was made) is described from the Pontic 
Range (Turkey). 1 have not seen the type of Koch and, therefore, refrain 
from accepting this name. Boissier united several species (for example, 
Scorzonera alpigena (C. Koch) Grossh., S. pulvinata Lipsch., S. 
subintegra (Boiss.) Thiebaut, etc.) under the name S. jacquiniana (C. 



47 

Koch) Boiss., with varieties alpina Boiss. and subintegra Boiss. (Fl. 
orient. Ill (1875) 757). As a result, S. jacquiniana sensu Boiss. is an 
aggregate unit. — The prostrate dwarf form of S. meyeri (C. Koch) Lipsch. 
is 1-3 cm high — f. nana (Grossh.). Askerova {Fl. Azerb. VIII (1961) 
519) described var. kubensis Asker. This variety, described from Tfandag 
(the Kuba Mountains of the Greater Caucasus), is characterized by the 
author as "forming a perennial plant with a many-headed root, (stem — 
S. L.) up to 5 cm high. Leaves pinnatisect, with short oblong-lanceolate 
segments, having dense, and short pubescent. Involucral bracts appressed 
gray-tomentose." I did not see specimens of this variety; therefore, this 
taxon remains unknown to me. 

11. S. kirpicznikovii Lipsch. sp. nova in Addenda XXVIII, 719. — 
Isaev in Fl. Azerb. VIII (1961) 520 (descr. ross.). — Podospermum canum 
C.A.M. Verzeichn. (1831) 62, p. p. quoad plant, talyschens.; DC. Prodr. 
VII, 1, 110, p. p. (excl. var. (3. glabratum and plant, talyschens.). — P. 
tenuisectum Grossh. and Sosn. in Zam. po Sist. i Geogr. Rast. Tbil. 13 
(1947) 67, quoad pi. talyschens. 

Perennial. Plant whitish-gray; root collar covered with remnants of 
sheaths of basal leaves; sheaths hard, shining, often incised. Stem scape- 
like, 6-12(15) cm high, simple, less often branched above and then 
with two capitula, pubescent, leafless or with isolated leaves, two times 
or one-third as long as basal leaves. Leaves predominantly basal, 
48 numerous, grayish-pubescent, up to 8 cm long, all leaves pinnatisect 
with large number of lateral lineral narrow (1-1.5 mm wide) segments, 
terminal segment oblong, narrowly linear; cauline leaves reduced, two 
or three, or absent. Capitula up to 1.5 cm long at flowering (including 
ligulate flowers), 1.8-2 cm long at fruiting. Involucre more or less 
pubescent, pubescence sometimes rubbed off; outer bracts smaller, 
triangular or triangular-lanceolate, often with cornicles at apex; inner 
bracts oblong-lanceolate, two to two and one-half times as long as outer 
ones. Ligulate florets grayish-yellow, pink on drying. Achenes slender, 
terete, 7-8(10) mm long, glabrous, smooth, with more or less distinct 
longitudinal ribs; stalk short; pappus dirty white, its bristles up to two- 
thirds plumose, scabrous above. 

Subalpine meadows. — Caucasus: Talysh. Endemic. Described from 
Talysh. Type in Leningrad. 

Note. The Talysh specimens of Meyer, treated by him as 
Podospermum canum, are sharply distinct from the Baku specimens, 
designated by me as the type of S. cana, by the structure of the achenes; 
they belong to the montane members of the subgenus Podospermum. 

12. S. grigoraschvilii (Sosn.) Lipsch. comb. nova. — Podospermum 
grigoraschvilii Sosn. in Dokl. Akad. Azerb. SSR, IV, 5 (1948) 212; 



48 

Grossh. Opred. Rast. Kavk. 508; Papava in Fl. Gruzii VIII, 638.— Ic: 
Fl. Gruzii VIII, Plate 433. 

Perennial. Plant completely glabrous, forming tussocks; root vertical, 
thick, many-headed, producing sterile rosette and flowering shoots. Stems 
ribbed-sulcate, short, 4—12 cm (at fruiting up to 25 cm) high. Leaves 
crowded at stem base; basal leaves up to 9 cm long, outer ' [lower] 
undivided, oblong-spatulate, narrowed or slightly broadened at apex, at 
base with two teeth; upper leaves oblong, pinnatipartite, terminal 
segments much larger than lateral ones, oblong-lanceolate or oblong- 
spatulate, obtuse; lateral segments short, linear; all [basal] leaves with 
petioles broadened at base, covering root collar; cauline leaves few (one 
to three), linear-spatulate, obtuse. Capitula at fruiting obconical, 1-1.5 cm 
wide. Involucre somewhat pubescent; bracts blackish, oblong-lanceolate, 
obtuse; inner bracts sometimes carinate. Achenes 0.8-1 cm long, olive- 
green slightly curved, with longitudinal ribs, glabrous; stalk indistinct; 
pappus white, almost two times as long as achene, pappus bristles fragile, 
up to two-thirds plumose, barbed above. Flowering August to September. 

Alpine areas in meadows. — Caucasus: Eastern Transcaucasia 
(Khevsuretia). Endemic. Described from Khevsuretia. Type in Tbilisi. 

Subgenus II. Pseudopodospermum (Lipsch. and Krasch.) Lipsch. 
comb. n. — Sect. Pseudopodospermum Lipsch. and Krasch. in Lipsch. 
F'-agm. Monogr. Scorzonera, I (1935) 70. — Species mostly with 
49 underground tubers. Ligulate florets yellow or red. Achenes stalked, more 
or less with tubercles or spinules along ribs. Leaves simple, linear or 
lanceolate. Involucral bracts not comiculate. By the nature of the achenes 
(presence of stalks), it approaches the species separated as members of 
the genus Podospermum DC. (i.e., subgenus Podospermum (DC.) 
Lipsch.), from which it differs in having simple, nonpinnate leaves and 
in most cases underground tubers. It differs from the members of 
subgenus Scorzonera by the presence of stalked achenes. 

Type of subgenus: S. millis M.B. 

Series 7. Molles Lipsch. in Addenda XXVIII, 719. — Root tuberous, 
near soil surface or deeper. Stems more or less tall, sulcate, leafy to 
apex. Ligulate florets yellow. 

13. S. mollis M.B. Fl. taur-cauc. Ill (1819) 522; DC. Prodr. VII, 1, 
122 (excl. var. leptophylla DC.) Stev. in Bull. Soc. Nat. Mosc. XXIX, 
4, 405; Boiss. Fl. or. Ill, 761 (quoad pi. tauric); Grossh. Fl. Kavk. IV 
(1934) 239; Lipsch. Fragm. Monogr. Scorzonera I (1935) 74; Krasch. in 
Fl. Yugo-Vost. VI, 447; Grossh. Opred. Rast. Kavk. 510.— 5. tomentosa 
auct. non L.: Pall. Reise III (1776) 513, 537, 654, 682, secund. Ldb.— 
Podospermum villosum auct. non M.B.: Stev. in DC. Prodr. VII, 1 (1838) 



49 

111.— P. molle Fisch. and Mey. in Ldb. Fl. ross. II, 2 (1845-1846) 
783 quoad pi. Ross, austral., taur. et ciscauc. — Scorzonera mollis 
var. caulescens O. Kntze. in Tr. Peterb. Bot. Sada X, 1 (1887) 
205. — Podospermum molle C.A.M. ex Stank, in Stank, and Tal. Opred. 
Vyssh. Rast. (1949) 685.— Ic: Bot. Mag. (1830) t. 3027; Lipsch. 
op. cit. (1935) 26; Fl. Yugo-Vost. VI (1936) 716.— Exs.: Callier, Herb, 
norm. edit, ab Dorfler, No. 4164; Lang and Szovits, Herb. Fl. ruth. No. 
77. 

Perennial. Arachnoid-canescent, with oval, thick underground tubers; 
root collar covered with sheaths, sometimes with numerous, somewhat 
violent, scarious scales. Stems solitary, (7)10-25(35) cm high, more or 
less pubescent, simple or branched from base, mostly densely leafy 
throughout, particularly in lower half. Leaves canescent, linear, hard, 
flat or folded, sometimes slightly undulate, broadened at base, drooping 
or somewhat arcuately bent, usually with wide midrib; cauline leaves 
sessile, somewhat amplexicaul, sometimes imbricately overlapping at 
base of stem. Capitula (1)3-7(12), cylindrical, (2)2.5-3.5 cm long, 
sometimes on peduncle somewhat thick at base. Involucre pubescent, 
becoming glabrescent; bracts few, unequal, more or less acuminate; outer 
bracts small, lanceolate; inner ones oblong-lanceolate, two to two and 
one-half times as long as outer. Ligulate florets yellow, exceeding 
involucre. Achenes narrowly cylindrical, stalked; outer achenes 
tuberculate along longitudinal ribs, scabrous; inner ones weakly 
50 tuberculate or smooth; pappus white, soft; pappus bristles plumose, five 
of them longer, barbed above. Flowering April to June. 

Steppes and stony, limestone, gypsum, clayey, and other dry 
slopes. — European Part: Bessarabia, Black Sea Region, Crimea, Lower 
Don, Lower Volga; Caucasus: Ciscaucasia. General distribution: Balkans- 
Asia Minor. Described from Crimea. Type in Leningrad. 

Note. The specimens found from time to time with inconspicuous, 
scape-like stems correspond to var. acaulis O. Kntze. (Tr. Peterb. Bot. 
Sada, X, 1 (1887) 205); they give the impression of having been grazed 
by cattle and abnormally developed. In the Askania-Nova steppes, var. 
virgata Lipsch. and Krasch. (Lipsch. Fragm. Monogr. Scorzonera I (1935) 
75) differs in having thick, distinctly sulcate, 40^5 cm high and 5-7 
mm thick stems, numerous virgate, simple, erect, slender branches, wider 
(6-10 mm wide) leaves with cartilaginous margins, and numerous (12- 
20 on single plant) capitula. 

Series 8. Leptophyllae Lipsch. in Addenda XXVIII, 719. — Stems 
scape-like, short, leafy only in lower part, less often with solitary leaves, 
never densely leafy. Tubers nut-shaped, oval or fusiform, at soil surface 
or deeper. Ligulate florets yellow or red. 



50 




51 Plate m. 

1 — Scorzonera leptophylla (DC.) Krasch. and. Lipsch.; 2 — 5. albicaulis Bge. 



51 

14. S. leptophylla (DC.) Krasch. and Lipsch. in Lipsch. Fragm. 
Monogr. Scorzonera I (1935) 78; Grossh. Fl. Kavk. IV (1934) 239; 
Grossh. Opred. Rast. Kavk. 510; Isaev in Fl. Azerb. VIII, 520.— 5. mollis 
var. leptophylla DC. Prodr. VII, 1 (1838) 122.— 5. mollis auct. non M.B.: 
Fisch. and Mey. PI. Szow. ined. p. p. — 5. szowitsii auct. fl. Cauc, pro 
max. parte. — S. undulata auct. non Vahl: Grossh. in Bot. Shorn. Azerh: 
GNU, I (1932) 53. — S. turcomanica auct. non Krasch, and Lipsch.: 
Grossh. Opred. Rast. Kavk. 510. 

Perennial. Plant (3)4-6(10) cm high at flowering, 15-25 cm at 
fruiting, with nut-shaped tuber borne below root collar near soil surface; 
roots fibrous, arising from tuber base; root collar covered with remnants 
of leaf sheath. Stems scape-like, one to three (sometimes up to seven), 
more or less tomentose, becoming glabrescent, slightly bent, leafy mainly 
in lower part. Leaves usually longer than stem at flowering, later on at 
fruiting, much shorter, narrowly linear, (1)2-3 mm wide, acuminate, 
glaucous or green, mostly pubescent, bent, usually folded lengthwise, 
52 less often flat, with cartilaginous margin, with more or less curly hairs. 
Capitula (one) three to seven, 2-2.5 cm long at flowering, with ligulate 
florets, 3.5-4 cm long at fruiting, cylindrical. Involucre pubescent, 
becoming glabrescent; outer bracts ovate-lanceolate, obtuse, inner bracts 
lanceolate or oblong-lanceolate, longer than outer, usually acuminate; 
ligulate florets yellow. Achenes stalked, narrowly terete, 14-15(18) nmi 
long, with longitudinal ribs; ribs verrucose, sometimes smooth; pappus 
grayish, pappus bristles plumose, five of them stiff, long, barbed above, 
scabrous. Flowering May to June (Plate III, Fig. 1). 

Stony-gravelly mountain slopes. — Caucasus: Eastern and Southern 
Transcaucasia. General distribution: Iran Afghanistan (northern). 
Described from Iran (vicinity of Deliman). Type in Leningrad. 

15. S. suberosa C. Koch in Linnaea XVII (1843) 275; Ldb. Fl. 
Ross. II, 2, 790; Boiss. Fl. or. Ill (1875) 761; Grossh. Fl. Kavk. 
IV (1934) 237; Lipsch. Fragm. Monogr. Scorzonera I, 80; Grossh. 
Opred. Rast. Kavk. 509. — S. aucheri Boiss. and Huet in Boiss. Diagn. 
pi. or. ser. II, 3 (1856) 92 sec. Boiss. (1875), sp. non vidi. — 5. tuberosa 
auct.: Walp. Repert. bot. system. VI (1846-1847) 351, non Pall.— 
Podospermum suberosum (C. Koch) Lipsch. in Fedde, Repert. XXXIII 
(1933) 168. 

Perennial. Resembling previous species; root thickened into oval or 
oblong tuber near soil surface. Stems (one) two or three (six), scape- 
like, (6)10-15(28) cm long, pubescent, leafy in lower part, leafless or 
with isolated reduced leaves above. Leaves linear, pubescent, longer or 
shorter than stem, flat or folded lengthwise, bent, along margin slightly 
undulate-crimped, with many veins, midrib mostly wide, white. Capitula 



52 

cylindrical (2)2.5-3.5 cm long; involucre pubescent, bracts somewhat 
scarious along margin, somewhat obtuse or slightly acuminate; outer 
bracts ovate-lanceolate; inner bracts lanceolate, longer than outer ones. 
Ligulate florets red, violet when dry. Achenes stalked, cylindrical, 15- 
18 mm long, with longitudinal ribs; outer achenes tuberculate-toothed 
along ribs, scarbrous; inner achenes more or less smooth; pappus grayish, 
pappus bristles plumose, five of them longer, in upper part barbed- 
scabrous. Flowering May to June. 

Stony-gravelly slopes. — Caucasus: ?Eastem Transcaucasia; Southern 
Transcaucasia. General distribution: Armenia and Kurdistan. Described 
from Somkhetia. Type in Berlin. 

Note. From all other species of the subgenus Pseudopodospermum 
found in the USSR, it differs by red (violet in dry condition) flowers. 

16. S. turkeviczii Krasch. and Lipsch. in Lipsch. Fragm. Monogr. 
Scorzonera I (1935) 83.— Ic: Lipsch. Ibid. (1935) Plates 30, 31. 

Perennial. Plant tomentose, arachnoid at some places, with root 
thickened into fusiform tuber; root collar more or less covered with leaf 
53 sheaths and scales. Stems one to three, scape-like, (4)6.5-12 cm high, 
pubescent, leafy in lower part. Leaves in type specimen bright green, 
with many (seven to nine) veins (midrib prominent, white), cartilaginous 
along margin, more or less distinctly undulate-crimped; basal leaves 
more or less broadly lanceolate, (4)6-12(16) mm wide, flat or folded, 
with obtuse callous cusp at apex, and sheathing base surrounding root 
collar; cauline leaves in lower part of scape-like stem, few, mostly 
narrowly lanceolate (4-6 mm wide), occasionally broadly lanceolate (10- 
15 mm wide), sessile, weakly amplexicaul. Capitula cylindrical, 1-1.2 
cm wide. Involucre pubescent, of a few bracts scarious along margin; 
outer involucral bracts ovate, subobtuse; inner ones oblong-lanceolate, 
. subobtuse; ligulate florets yellow. Achenes unknown. Flowering May to 
June. 

Stony, clayey, and limestone slopes. — Caucasus: Ciscaucasia 
(Krasnodar Territory). General distribution: eastern Anatolia. Described 
from former Kars Region. Type in Leningrad. 

Note. Our plants belong to a separate variety — var. kubanica Krasch. 
and Lipsch., op. cit. 

17. S. turcomanica Krasch. and Lipsch. in Lipsch. Fragm. Monogr. 
Scorzonera I (1935) 80. — S. mollis auct. fl. turcom. non M.B. — S. 
tuberosa auct. quoad pi. turcom. p. p. — S. raddeana auct non Winkl.; 
Lipsch. in Fl. Turkm. VII (1960) 304, p. p. — IS. stenophylla Boiss. and 
Hausskn. ex Lipsch. op. cit. (1935) 80. — Ic: Lipsch. op. cit. (1935) 
Plate 29; Lipsch. op. cit. (1960) Plate XL VI, Fig. 1. 



53 

Perennial. Whole plant farinose; root gradually thickened into tuber, 
sometimes spherical; root collar densely covered with sheaths of basal 
leaves, partly incised but not fibrous. Stems numerous, scape-like, 10-20 
cm high, leafy only in lower part. Leaves filiform, narrowly linear, 1-2 
mm wide, bent or drooping, farinose, flat or rolled, entire, with callose 
thickening at apex, acuminate, mostly longer than often leafy stem. 
Capitula 6-10(19), mostly narrowly cylindrical, (2)2.5-3.5 cm long. 
Involucre tomentose, becoming glabrescent; outer bracts ovate-lanceolate; 
inner lanceolate, two to two and one-half times as long as outer; all bracts 
somewhat scarious on margin, obtuse. Ligulate florets yellow. Achenes 
on small stalk, cylindrical, with longitudinal ribs, acerate along ribs; pappus 
dirty gray or blackish, its bristles plumose to three quarters length, barbed 
above. Flowering April to May. 

Steppes and stony and marly slopes. — Soviet Central Asia: 
mountainous Turkmenia. General distribution: Iran (northern), 
Afghanistan. Described from Kopetdag. Type in Leningrad. 

Note. See note on S. raddeana Winkl. 

54 18. S. raddeana Winkl. in Tr. Peterb. Bot. Sada XI, 1 (1890) 150; 

Lipsch. in Fl. Turkm. VII, 304, p. p.— Ic: Winkl. op. cit. t. Ill, fig. 1, a; 
Lipsch. op. cit. Plate XL VI. 

Perennial. Plant slightly arachnoid and canescent; root apparently 
thickened into tuber; root collar densely covered with brownish sheaths 
of basal leaves. Stem 25 cm high, in form of leafy sulcate scapes, branched 
almost from base. Leaves linear, up to 5 cm wide, acuminate; basal leaves 
broadened into sheaths; cauline semiamplexicaul; all leaves along margin 
weakly undulate-crimped; peduncles slightly exceeding leaves. Capitula 
large (3.5 cm long and 1.5 cm wide in fruits), solitary, terminal on 
branches. Involucre arachnoid pubescent; bracts unequal, margin white- 
membranous. Ligulate florets red, on drying lilac-colored (needs to be 
confirmed in nature). Achenes terete, brown, ribbed, densely spiny, without 
swollen stalks; pappus dirty gray or blackish, almost as long as achene; 
pappus bristles stiff, plumose almost to three quarters of their length, in 
upper part toothed-scabrous. Flowering April to May. 

Gravelly desert. — Soviet Central Asia: Mountainous Turkmenia 
(between Khodzha-Kala and Bami). Endemic. Described from the place 
indicated above. Type in Leningrad. 

Note. Only one specimen of this species is known — the authentic 
specimen; the species has never been collected subsequently. It is quite 
close to S. turcomanica Lipsch. and Krasch., from which it is distinguished 
by having broader leaves, a scapelike but leafy stem that is branched 
from the very base, red (needs verification in nature from living material) 
ligulate florets, and achenes without stalks. In the Flora of Turkmenia 



54 

(VII, p. 304), I united S. raddeana C. Winkl. with S. turcomanica Krasch. 
and Lipsch. into a single species, under the first (priorable) name. 
However, this question cannot be completely resolved. Therefore, in the 
present work, with the aim of drawing attention to both taxa, they are 
treated as separate entities, which are presented in the identification key 
and in the text. Compare my note in the Flora of Turkmenia (p. 306). — 
This species is included tentatively under the subgenus 
Pseudopodospermum. Although it does not have a stalked achene, it is 
close to the members of the subgenus in other characters. 

Subgenus III. Scorzonera. — Euscorzonera Rouy, Fl. Fr. X (1908) 9; 
Consp. Fl. Fr. (1927) 169; Lipsch. Fragm. Monogr. Scorzonera II (1939) 
9. — Subgen. Euscorzonera Lipsch. Fragm. Monogr. Scorzonera I (1935) 
7.— Gen. Scorzonera Cass, in Diet. sc. nat. XXV (1822) 63, 264; Rchb. 
Fl. Germ. exc. (1830) 275; Endlich. Gen. pi. (1838) 497; Benth. and 
Hook. f. Gen. pi. II (1873) 531 (excl. sect. Pterachaenia and 
Podospermum); Boiss. H. or. Ill (1875) 755-783, excl. sect. Podospermum 
(pp. 757-760) and Epilasia (pp. 781-783); O. Hoffm. in Pflanzenfam. 
IV, 5 (1893) 365, excl. sect.: Pterachaenia Benth., Podospermum DC. 
and Epilasia Bge. — Achyroseris Sch. Bip. in Nov. Act. Acad. Caesar. 
Leopol.-Car. Natur. Cur. XXI, 1, (1845) 165. — Scorzonerae sect, la Less. 
Synops. Compos. (1832) 134. — Sect. Euscorzonera DC. Prodr. VII, 1 
(1838) 117; Ldb. Fl. Ross. II, 2 (1845-1846) 790, p. p.; Boiss. Fl. or. Ill 
(1875) 760-776, p. p. — Fleischeria Hochst. and Steud. ex Endlich. Gen. 
55 pi. (1838) 491.— Gelasia Cass, in Bull. Soc. Philom. Paris (1818) 33 
(opusc. non vidi); in Diet. sc. nat., XVIII (1820) 285; XLII (1826) 81; 
Less. Synops. Compos. (1832) 134 (sub nom. Galasia); W. Koch. Synops. 
Fl. Germ, et Helv. (1837) 426, pro genere; DC. Prodr. VII, 1 (1838) 123, 
pro sect.; Endlich. Gen. pi. (1838) 497, pro gen. Galasia. — Lasiospora 
Cass, in Diet. sc. nat. XXV (1822) 306; Less. Synops. Compos. (1832) 
134; DC. Prodr. VII, 1 (1838) 124, pro sect; Ldb. Fl. Ross. II, 1 (1845- 
1846) 796, pro sect.; Sch. Bip. op. cit. (1845) 166, pro gen; Boiss. Fl. or. 
Ill (1875) 776-781, pro sect. — Lasiospermum Fisch. Catal. Hort. Gorenk. 
(1812) 34, non Lag.— Poly clada DC. Prodr. VII, 1 (1838) 125, pro sect. 

Achenes without hollow swollen stalks at base. All leaves simple, 
entire, sometimes pinnatifid only in S. bicolor. Involucral bracts not 
comiculate at apex. Achenes glabrous or lanate. 

Type of subgenus: type of genus. 

Section \. Incisae Lipsch. in Lipsch. Fragm. Monogr. Scorzonera 1 
(1935) 91. — Plant with pinnatifid or pinnatisect leaves. Leaves quite 
variable in degree of cutting, in several species with serrate margin. 



55 

Flowers bicolored: ligule yellow, blackish-purple near base. Achenes 
without stalks, white or flesh-colored, ribbed (ribs corky), sulcate. 

19. S. bicolor Freyn and Sint. in Oesterr. Bot. Zeitschr. XLIII (1892) 
266; Lipsch-Fragm. Monogr. Scorzonera I (1935) 102; Grossh. Opred. 
Rast. Kavk. 510; Isaev in Fl. Azerb. VII, 521. — S. incisa var. dentata 
Bordz. in Zap. Kiev. Obshch. Estestv. XXV, 1 (1915) 131; Grossh. Fl. 
Kavk. IV (1934) 238.-5. karjaginii Grossh. in Bot. Shorn. Azerb. GNU 

I (1932) 52, descr. ross.; Tr. Azerb. Otd. Zakavk. Fil. Akad. Nauk SSSR, 
Sekt. Bot. I (1933) 62, descr. latin; Fl. Kavk. IV (1934) 238.— Ic: Lipsch. 
op. cit. (1935) Plates 38, 39. 

Perennial. Plant arachnoid-pubescent, later glabrescent, with vertical 
root; root collars covered with numerous hard, glossy, partly torn sheaths. 
Stems 30-50 cm high, pubescent, later glabrescent, erect or somewhat 
ascending, solitary or a few (three to five), sulcate, leafy, branched in 
upper part. Leaves hard, pubescent, with distinct midrib, more or less 
carinate beneath, margin cartilaginous, variously incised; basal leaves 
numerous, narrowed into long sulcate petiole, oblong-lanceolate, lamina 
15-20 cm long and 3 cm wide, sharp-toothed, pinnatifid or less often 
entire; upper cauline leaves highly reduced, sessile, amplexicaul, oblong- 
lanceolate or lanceolate, toothed or incised, with obtuse auricles at base. 
Capitula large, on long peduncles, 2-2.5 cm long and 0.8-1 cm wide at 
flowering, up to 4 cm long and 2 cm wide at fruiting. Involucre of 
imbricate, glabrous, hard bracts more or less scarious along margin; outer 
56 bracts ovate-deltoid; inner ones oblong-lanceolate, two to four times as 
long as outer ones. Florets and ligules yellow, corolla throat black-purple. 
Outer achenes 15-20 mm long, white or flesh-colored, with corky ribs, 
deeply sulcate, with obtuse teeth along ribs, basally fused outer achenes 
often not filled, flattened; pappus dirty white, 18(25) mm long, pappus 
bristles stiff, plumose, barbed above. Flowering June to August. 

Stony gravelly slopes. — Caucasus: Southern Transcaucasia. General 
distribution: Armenia and Kurdistan. Described from Turkey. Type in 
Freyn' s herbarium. 

Section 2. Purpureae Lipsch. in Lipsch. Fragm. Monogr. Scorzonera 

II (1939) 104. — Root collar densely hairy. Stem mostly up to 80 cm 
high, simple or branched, with one to five capitula, usually densely leafy. 
Achenes glabrous, somewhat swollen at base. Inner involucral bracts 
subobtuse, distinctly white-membranous along margin. Florets pink, red 
or violet. 

20. S. purpurea L. Sp. pi. (1753) 791; Willd. Sp. pi. Ill, 3 (1803) 
1502, descr. emend.; M.B. Fl. taur.-cauc. II, 236; DC. Prodr. VII, 1, 118; 



56 

Ldb. Fl. Ross. II, 2, 791, p. max. p.; Boiss. Fl. or. Ill, 768; Schmalh. Fl. 
II, 142; Kryl. Fl. Alt. Ill, 731; Syreistsch. 111. Fl. Mosk. Gub. Ill, 328; 
Hegi, 111. FL, IV, 2 (1929) 1054; Krasch. in Fl. Yugo-Vost. VI, 452; 
Lipsch. Fragm. Monogr. Scorzonera II, 106; Kryl. Fl. Zap. Sib. XI, 
2987. — S. purpurea subsp. eupurpurea Herrai. in Hegi, op. cjt. (1929) 
1054. — Podospermum purpureum Koch ex Steud. Nomencl. bot. ed. 2, 
II (1841) 546.— Ic: Rchb. Ic. Fl. Germ. XIX, t. 1382; Syreistsch. op. 
cit. 328; Hegi, op. cit. t. 276, fig. 3; fig. 737, 738, 746. Exs.: GRF No. 
1622. 

Perennial. Root vertical, cylindrical, root collar densely setose with 
remnants of petioles. Stems (8)15^5(80) cm high, solitary or a few, 
glabrous, arachnoid at isolated places, especially below capitula or at leaf 
base, simple (and then with single capitulum) or branched above, with 
two to five solitary terminal capitula. Leaves narrowly linear, herbaceous, 
1-3(4) mm wide, entire, triangular-grooved, glabrous or weakly arachnoid- 
pubescent; basal leaves 5-25 cm long, somewhat narrowed at base, with 
more or less densely pubescent sheaths; cauline leaves usually many (five 
to seven), sessile, somewhat broadened at base. Capitula medium-sized; 
up to 2.5 cm long and 0.4-1 cm wide. Involucre oblong-cyhndrical, almost 
glabrous or slightly pubescent mainly in lower part; outer involucral bracts 
58 ovate, often with brown border along margin; inner bracts two times as 
long as outer ones, margin scarious, lanceolate, obtuse. Ligulate florets 
light purple or violet-pink, with vanilla flavor, one and one-half to two 
times as long as involucre. Achenes up to 12 mm long, glabrous, with 
ring of hairs only below pappus, ribbed, ribs smooth; pappus bristles 
plumose. Flowering May to June. (Plate IV, Fig. 1.) 

Steppes, wet and steppe meadows, grassy slopes, sands, edges of 
steppe pine forests, gravel and limestone slopes. — European Part: Black 
Sea Region, Middle Dnieper, Baltic Region, Volga-Don, Upper Volga, 
Volga-Kama, Trans- Volga, Lower Don; Western Siberia: Upper Tobol, 
Irtysh; Soviet Central Asia: Aralo-Caspian, Balkhash Region. General 
distribution: central Europe, Mediterranean. Balkans-Asia Minor. Described 
from central Europe. Type in London. 

21. S. rosea Waldst. and Kit. Descr. pi. rar. Hung. II (1805) 127; 
Willd. Sp. pi. Ill, 3, 1502; DC. Prodr. VII, 1, 118; Boiss. Fl. or. Ill, 769, 
p. min. p.; Hegi, 111. Fl. VI, 2 (1929) 1055; Lipsch. Fragm. Monogr. 
Scorzonera II, 108. — S. purpurea p. rosea Ldb. Fl. Ross. II, 2 (1845- 
1846) 792. — S. purpurea ssp. rosea (Waldst. and Kit.) Herrm. in Hegi, 
op. cit. (1929) 1055.— Ic: Waldst. and Kit. op. cit. (1805) t. 121; Rchb. 
Ic. fl. Germ. XIX, 1382; Hegi, op. cit. (1929) 1056, fig. 739; Javorka 
and Csapody, Magyar Fl. Kepekben, i. 556, No. 4000 (nigra), t. XL 
(color). 



57 




57 



Plate IV. 
1 — Scorzonera purpurea L.; 2 — S. crispa M.B. 



58 

Perennial. Root fusiform-cylindrical; root collar covered with 
numerous blackish-brown hairs. Stems 15-60 cm high, erect or ascending, 
simple (very rarely branched near base and then with single branch) with 
single capitulum. Leaves lanceolate to linear-lanceolate, 3.5-4 mm wide, 
flat, glabrous; basal leaves often as long as stem; cauline leaves much 
shorter. Capitula large, up to 5 cm long in full bloom; involucre oblong- 
cylindrical, 2-3 cm long, weakly lanate at base; involucral bracts 16-20; 
outer bracts deltoid, oblong, obtuse, scarious along margin; inner bracts 
two to three times as long as outer, lanceolate, obtuse, mostly with brown 
spot. Florets pale-pinkish, peripheral ones two times as long as involucre. 
Achenes 15 mm long, pale yellow or pale reddish-brown, ribbed, ribs in 
upper part toothed-scabrous; pappus plumose. Flowering June to July. 

Alpine and subalpine meadows, open woodlands and scrubs, 
limestones. — European Part: Upper Dniester (Carpathians). General 
distribution: central Europe, Mediterranean, Balkans-Asia Minor. Described 
from Hungary. Type in Budapest; isotype in Leningrad. 

Note. This is a montane race giving rise to a series of transitions to 
S. purpurea. The type specimen of S. rosea differs from S. purpurea by 
having wider, flat leaves, larger capitula, a larger number of involucral 
bracts and ribs on the achenes, these being toothed-scabrous in the upper 
part. In the Carpathians, it is a typical plant of the subalpine meadows, 
also extending into the alpine zone. 

59 Section 3. Egregiae (Kult.) Lipsch. Comb. n. — § (subsect.) Egregia 

Kult. Tau-Sagyg i E'kol. Osnovy Vvedeniya Ego v Kulture* (1938), 
108. — Subgen. Egregia (Kult.) Ovcz. in Soobshch. Tadzh. Fil. Akad. Nauk 
SSSR XX (1949) 54. — Semishrubs, forming hemispherical tussocks. 
Nonspiny. Elastic latex fibers distinctly visible on breaking underground 
organs. Stems (caudices) never hard- woody; current year's stems not 
producing rosettes, simple or less often weakly branched. Achenes terete, 
ribbed, along ribs more or less tuberculate-scabrous, pubescent. Pappus 
white, pappus bristles plumose, inner bristles longer, barbed above. 
Capitula many-flowered. 

22. S. tau-saghyz Lipsch. and Bosse. in Tr. N.-I. Labor. Tresta 
"Kauchuknos," 4 (1930) 27; Tr. N.-L Inst. Promyshl. 465 (1931) 5; Lipsch. 
sistem. Zam. ob Aziat. Skorts. (1932) 11; Lipsch. Fragm. Monogr. 
Scorzonera I (1935) 18; Kult. Tau-Sagyz i E'kol. Osnovy Vvedeniya Ego 
V Kul'ture (1938) 61.— S. vavilovii Kult. in Kult. op. cit. (1938) 56.— 
S. mariae Kult. in Kult. op. cit. (1938) 59. — S. karataviensis Kult. in 



*Tau-Sagyz and the Ecological Basis of Its Introduction into Cultivation — Translator. 



59 

Kult. op. cit. (1938) 66.-5. longipes Kult. in Kult. op. cit. (1938) 70.— 
5. kirghisorum Afan. in Tr. Tadzh. Bazy Akad. Nauk SSSR, VIII, 1938 
(1940) 474. — S. tadshikistanica Ovcz. in Soobshch. Tadzh. Fil. Akad. 
Nauk SSSR XII (1949) 8, nomen.— 5. rindak Ovcz. in Soobshch. Tadzh. 
Fil. Akad. Nauk SSSR XX (1949) 53 in adnot.— 5. usbekistanica 
Czevrenidi and Bondarenko in Dokl. Akad. Nauk Uzb. SSR 3 (1950) 37, 
nomen; Galkina in Tr. Inst. Bot. Akad. Nauk Uzb. SSR I (1932) 5-23 
(detailed description of species in Russian); Kovalevskaja in PL Uzb. VI 
(1962) 436 and Appendix, 516 (descr. lat.). — Ic: Lipsch. and Bosse, op. 
cit. (1931) Plates 1 and 2 (color); Kult. op. cit. (1938) Figs. 1, 29, 39, 
44, 49, 55, 60, 73; Fl. Uzb. VI (1962) Plate XLIV, Fig. 2. 

Perennial. Semishrub forming loose or somewhat compact 
hemispherical tussocks; whole plant more or less sericeous with simple 
flexuous hairs, glabrescent; but sometimes bases of fallen hairs visible. 
Rhizome strong, deeply embedded in soil, strongly branched above, many- 
headed, forming tussock of intertwined underground stems, terminating 
above soil surface in rosettes of more or less numerous (6-25) basal 
leaves and erect, somewhat flexuous, 3 to 165 flowering shoots. Thin 
elastic latex fibers distinctly visible in laticifers on breaking roots and 
underground stems; latex milky-white or yellowish-green; aerial stems 
densely covered with numerous imbricate, grayish-brown, weakly glossy, 
rigid, basally broadened into deltoid, fibrous sheaths of dried and partly 
torn leaves of previous years. Leaves of rosette (6-25) broadened at base 
into sheaths, grayish-green, variable, narrowly linear, with reddish-brown 
subulate apex, often uncinate, distinctly triquetrous below, somewhat flat 
60 above, often incurved, carinate, or fistular, narrowed at both ends, roundish 
in cross section, or subulate-edged, carinate-subulate; all leaves coriaceous, 
three- to five- veined, with broad light yellow midrib beneath; sheaths not 
green, yellowish-brown, glossy, dorsally smooth or roundish-carinate, with 
dense tuft of simple multicellular hairs, initially white, later rusty; up to 
eight parallel reddish-brown veins visible at base of leaf sheath. Flowering 
shoots erect, developing every year (one to three) from base of leaf rosette, 
drying out and falling after fruiting, 5-40 cm long, with 7-23 leaves, 
longer than rosette leaves, sulcate, sometimes fistular, simple, with single 
capitulum, or branched, with two to four capitula; clavately thickened or 
not below capitula; cauline leaves alternate; all leaves somewhat broadened 
at base; lowermost leaves scarious, scaly, aristate-acuminate; middle 
cauline leaves mostly flat, sometimes carinate, somewhat divergent; upper 
leaves reduced, bent, filiform, hairy. Capitula solitary or in groups of a 
few, cylindrical, obconical or pyriform, (1.3)2.3-4(4.5) cm long; involucre 
many-rowed (three to six), with 15-70 bracts; involucral bracts imbricate, 
highly unequal in shape, size, pubescence, and color; outer and middle 



60 

bracts shorter, deltoid or lanceolate, more or less carinate, sometimes 
crestate, some bracts with callose, mildly serrate, tuberculate thickening 
in upper part of keel; inner bracts longest (up to 2.2 cm), oblong-lanceolate 
to linear, obtuse or somewhat acuminate, with scarious margin, mpstly 
without keel or with barely visible keel toward apex, or more or less 
pubescent. Ligulate florets more or less numerous (30-50 in capitulum), 
yellowish with dark red veins (turning red from outside on drying), longer 
than involucre; sparse tuft of white hairs in upper part of corolla tube on 
outside near base of ligule. Achenes terete, 6-14 nam long, with 10 ribs, 
ribs smooth or tuberculate-scabrous, sometimes with sharp spinules, usually 
densely pubescent; pappus snow-white, its bristles plumose, inner bristles 
4-6(10), longer and fragile, toothed above. Flowering May to July. 

Stony gravelly northern slopes and plateaus, clayey ridges (Karatau 
Range); slopes of adyrs (Fergana); steep north-facing slopes at 1,900- 
2,100 m (Turkestan Range). — Soviet Central Asia: Syr-Darya (Karatau 
Range); Pamiro-Alai (Fergana valley, Turkestan and Hissar ranges). 
Endemic. Described from Syr-Darya Karatau. Type in Moscow. 

Note. An unusually polymorphic species, extremely variable in the 
shape and width of leaves, consistency and number of leaves in the 
rosettes, size, shape, pubescence and color of the involucral bracts, nature 
61 of the pubescence and achene sculpture, structure of the pappus hairs, 
etc. The division of tau-sagyz into a large number of smaller species is 
not at all convincing to me, because the characters given as the basis for 
separating them, with the variety of ecological conditions in which tau- 
sagyz is found and its fragmented range are of an intermediate, unstable 
nature. Capable of regeneration. Tau-sagyz, a well-known rubber 
producing plant, has generated a voluminous literature in the USSR; it 
has been introduced into cultivation. 

The study of the specimens of Scorzonera acantholimon Hand.-Mazz., 
collected in Turkey and kindly sent to me by Davis from Edinburgh, 
showed that this species, which was earlier considered close to 5. tau- 
saghyz Lipsch. and Bosse, has nothing in conmion genetically with the 
latter. The types of the species described by M.V. Kultiasov are preserved 
in Leningrad. The type of S. kirghisorum Afan. is also there. The type of 
5. rindak Ovcz. is in Dushanbe, and that of S. uzbekistanica Czevr. and 
Bond, in Tashkent. 

Section 4. Piptopogon C.A.M. ex Turcz. in Bull. Soc. Nat. Mosc. 
XXI, 3 (1848) 97 (Fl. baic.-dahur.); Lipsch. Fragm. Monogr. Scorzonera 
II (1939) 66 (descr. emend.). — Gen. Achyroseris Sch. Bip. in Nov. Acta 
Acad. Caes. Leopol. -Carol. Nat. Cur. XXI (1845) 165.— Sect. 
Macrospermae Nakai in Report. Inst. Sc. Res. Manch. I, 6 (1937) 168, in 
adnot. — Capitula large, especially in fruiting plants, (3.5)5-8 cm long 



61 

and (1)1.5-2(3) cm wide. Achenes elongate, (2)4^.5 cm long, apex 
narrowed into a beaklike structure. Involucre many-rowed; outer involucral 
bracts reduced, ovate-deltoid; inner ones linear-lanceolate, usually rather 
long (up to 6 cm); pappus dirty-yellow, usually falling off entirely. Ligulate 
florets yellow and pink, resembling Tragopogon species in external 
appearance. 

Series 9. Macrospermae Lipsch. in Addenda XXVIII, 719. — Ligulate 
florets yellow. 

23. S. albicaulis Bge. in Mem. Acad. Sc. Petersb. div. sav. II (1835) 
114; DC. Prodr. VII, 1, 117; Maxim, in Bull. Acad. Sc. Petersb. XXXII, 
4 (1888) 495; Koni. and Alis. Opred. Rast. Dal'nevost. Kr. II, 1091; 
Lipsch. Fragm. Monogr. Scorzonera II, 75.^5. albicaulis Bge. f. rosea 
Nakai 2Xid flavescens Nakai in Report. Inst. Sc. Res. Manch. I, 6 (1937) 
169. — 5. macrosperma Turcz. in DC. Prodr. VII, 1 (1838) 121, descr.; in 
Bull. Soc. Nat. Mosc. XI, 1 (1838) 95, nomen. XXI, 3 (1848) 97 (Fl. 
baic.-dahur.) descr.; Ldb. Fl. Ross. II, 2, 795; Maxim. Prim. fl. amur. 
177; Rgl. Tent. Fl. Ussur. 302; Nakai, op. cit. (1937) 169.— Piptopogon 
macrospermus C.A.M. ex Turcz. in Bull. Soc. Nat. Mosc. XI, 1 (1838) 
95, nomen; Sch. Bip. in Nov. Acta Acad. Caesar. Leopol. -Carol. Nat. 
Cur. XXI (1845) 165. — Achyroseris macrosperma Sch. Bip. op. cit. (1845) 
166.—? S. radiata p. linearifolia DC. Prodr. VII, 1 (1838) 122.— Ic: 
Nakai op. cit. (1937) t. II and III. 
62 Perennial. Root collar covered with reddish-brown sheaths of previous 

year's leaves. Stems up to 1 m high, erect, fistular, sulcate, floccose, 
later almost glabrous. Leaves flat, linear or broadly linear, acuminate, 
five- to seven-veined, glabrous or weakly pubescent, serrulate under hand 
lens; basal leaves up to 0.4 m long, 0.7-1.8(2) cm wide; cauline leaves 
similar to basal leaves in shape and size, sessile, slightly broadened at 
base, amplexicaul, reduced in size above. Capitula in umbellate corymb 
(f. umbellata Lipsch.) or on lateral peduncles, cyhndrical, up to 4.5 cm 
long and 1-1.2 cm wide at flowering. Involucre many-rowed, arachnoid- 
pubescent or almost glabrous; outer bracts deltoid-ovate, smaller, middle 
obovate, somewhat acuminate; inner ones linear-lanceolate, somewhat 
scarious along margins, obtuse or somewhat acuminate, two times as long 
as middle ones. Ligulate florets yellow, pinkish when dry, almost one 
and one-half times as long as involucre. Achenes large, up to 2.5 cm 
long when mature, reddish-brown, gradually tapered toward apex to form 
beak, with somewhat sharp ribs; pappus dirty-yellow, deciduous, pappus 
bristles plumose, connate at base into ring, five of them long, 
sharp-pointed, barbed above. Flowering July to August (Plate III, 
Fig. 2.) 



62 




63 



Plate V. 
1 — Scorzonera seidlitzii Boiss.; 2 — 5. radiata Fisch. 



63 

Dry meadows, thin evergreen forests, scrub, and wastelands. — Eastern 
Siberia: Dauria; Far East: Zeya-Bureya, Ussuri, Uda Region. General 
distribution: northeastern China, Korean Peninsula. Described from 
northern China {S. albicaulis Bge.) and Nerchinsk Dauria (S. macrosperma 
Turcz.). Type in Leningrad. 

Series 10. Bracteosae Lipsch. in Addenda XXVIII, 720. — Capitula 
surrounded by more or less large adnate bracts. Leaves broadly lanceolate 
or ovate-lanceolate, 2.5^ cm wide. 

24. S. bracteosa Winkl. in Tr. Peterb. Bot. Sada XI, 2 (1890) 324; 
Lipsch. Fragm. Monogr. Scorzonera II (1939) 68; Kovalevskaja in Fl. 
Uzbek. VI, 438. — S. inula Lipsch. in Lipsch. Sistem. Zam. ob Aziat. 
Skorts. (1932) 19; Lipsch. in Fedde, Repert. XXXIII (1938) 166.— Ic: 
Lipsch. op. cit. (1933) 33, Fig. 9 (sub nom. S. inula); Lipsch. op. cit. 
(1939) Plate 11. 

Perennial. Plant (10)15-60 cm high, with one or many stems. Stems 
sulcate, more or less pubescent, glabrescent, usually with corymbose 
branching, sometimes sparsely leafy, and then intemodes long. Leaves 
pubescent or glabrescent, with many (up to seven) veins, hard, serrulate 
under hand lens, flat or slightly repand; basal leaves 2.5-3.0 cm long, 
64 narrowed into long petiole, with somewhat broadened sheath; cauline 
leaves gradually reduced above, alternate or almost opposite, broadly 
lanceolate, (1)2-4 cm wide, amplexicaul; bracteal leaves (1)2-7 cm long, 
appressed to base of capitulum, variable in number (one to four) and 
size, similar to upper cauline leaves (i.e. also ovate-lanceolate or less 
often lanceolate), sometimes exceeding capitula. Capitula (2)5-8(15), 5.5 
cm long at flowering, up to 8 cm long and up to 3.5 cm wide at fruiting. 
Involucre somewhat pubescent; outer bracts reduced, (0.3)0.5-0.6 cm long, 
deltoid-ovate, sometimes somewhat carinate; middle ones 2-3(3.5) cm 
long, oblong-lanceolate; inner ones 4-5(7) cm long, linear-lanceolate. 
Ligulate florets yellow. Achenes up to 4 cm long, brownish-red when 
mature, narrowed above into beak, scabrous on ribs or almost smooth; 
pappus dirty-yellow, deciduous, its bristles up to two-thirds plumose, 
barbed above. Flowering June to July. 

Stony steppe slopes, screes of conglomerates, etc. — Soviet Central 
Asia: Pamiro-Alai. Endemic. Described from Hissar. Type in Leningrad. 

25. S. alaica Lipsch. in Lipsch. Fragm. Monogr. Scorzonera II (1939) 
71. 

Similar to the preceding species in external appearance, shape and 
size of vegetative and reproductive organs. Distinguished by pink ligulate 
florets and reduced bracts. Flowering June. 



64 

Soviet Central Asia: Pamiro-Alai (Alai Valley, Gulcha). Endemic. 
Described from Alai. Type in Leningrad. 

Note. Described from three herbarium sheets collected by V.A. 
Tranzshel. The available material shows a clear difference between this 
Alai race and 5. bracteosa; however, additional field observations of this 
race are needed. 

Series 11. Tragopogonoideae Lipsch. in Addenda XXVIII, 720. — 
Capitula not surrounded by somewhat large adnate bracts, leaves linear 
or broadly linear, herbaceous, up to 2 cm wide. Ligulate florets pink. 

26. S. albertoregelia Winkl. in Tr. Peterb. Bot. Sada, XI, 2 (1890) 
322; Lipsch. Fragm. Monogr. Scorzonera II (1939) 68; Kovalevskaja in 
Fl. Uzb. VI, 437 (pro syn. 5. tragopogonoides Rgl. and Schmalh.). — Ic: 
Lipsch. in Byull. Mosk. Obshch. Isp. Prir., Otd. Biol., XLII, 2 (1933) 
155, Fig. 2; Lipsch. op. cit. (1939) Plate 10. 

Perennial. Dwarf plant, 2-3.5 cm high. Stems prostrate, numerous 
(up to 10). Leaves narrowly linear, 0.2 cm wide, glaucescent, glabrous, 
or somewhat pubescent, serrulate under hand lens, mostly three-veined. 
65 Capitula large, 4-4.5 cm long, one and one-half to two times as long as 
stem. Involucre arachnoid-tomentose; outer involucral bracts deltoid, 7.5- 
8 mm wide, with uncinate apex; inner bracts linear-lanceolate, 4 cm long, 
more or less broadly membranous along margin. Ligulate florets pink or 
light violet(?). Achenes 2.5 cm long, reddish-brown, narrowed above into 
beak; pappus dirty yellow, wholly deciduous, pappus bristles up to two- 
thirds plumose, barbed above, scabrous. Rowering August. 

Soviet Central Asia: Pamiro-Alai (Karategin). Endemic. Described 
from Karategin. Type in Leningrad. 

Note. The species was described by Winkler from a single specimen; 
it has never been collected again and requires additional study. Possibly, 
the most typical characters of S. albertoregelia — numerous dwarf stems 
and capitula exceeding the stems — indicate that the specimen used for 
describing this species was regenerated from a plant grazed by cattle 
{? Scorzonera tragopogonoides). 

27. S. tragopogonoides Rgl. and Schmalh. in Tr. Peterb. Bot. Sada, 
V (1877) 635; Lipsch. Sist. Zam. ob. Aziat. Skorts. (1932) 22; Lipsch. 
Fragm. Monogr. Scorzonera II (1939) 82; Popov in Tr. Uzb. Gos. Univ. 
Nov. Ser. 27, Biol. 14 (1941) 93; Lipsch. in Fl. Turkm. VII, 307; 
Kovalevskaja in Fl. Uzb. VI, 437, p. max. p. — S. glabra Winkl. in Tr. 
Peterb. Bot. Sada, XI, 2 (1890) 323.-5. korshinskyi Lipsch. op. cit. (1932) 
18; Lipsch. in Fedde, Repert. XXXIII (1933) 165.— 5. nikitini Lipsch. 



65 

op. cit. (1932) 20; Lipsch. op. cit. (1933) 166. — S. crassicaulis Rech. f. 
in Symb. Afghan. II (1955) 191.— Ic: Lipsch. op. cit. (1939) Plate 15, 
16; Rech. f. op. cit. (1955) fig. 133. 

Perennial. Plant 20-60 cm high, with fusiform root; root collar covered 
with remnants of previous year's leaves. Stems solitary, erect, sulcate, 
pubescent, glabrescent, more or less densely leafy. Leaves flat or folded, 
herbaceous, narrowly or broadly Unear, 0.1-0.8 cm wide, many-veined, 
serrulate under hand lens, scabrous; basal leaves somewhat broadened 
into sheaths; cauline leaves sessile, somewhat broadened at base, 
amplexicaul, all leaves acuminate. Capitula axillary, at variable height, 
not forming candelabrumlike corymbose inflorescence; peduncles clavately 
thickened below capitula, decorated with small membranous leaves. 
Capitula 5-5.5 cm at flowering, up to 6-6.5 cm long at fruiting. Involucre 
cylindrical, many-rowed, imbricate, with fine arachnoid pubescence; 
involucral bracts often with distinct carinate midrib; outer bracts small, 
deltoid or deltoid-ovate, usually acuminate, sometimes with falcate cusp; 
middle ones ovate-deltoid or lanceolate, almost two times as long as outer; 
inner ones Unear-lanceolate, membranous along margin, largest up to 6 
cm long (in fruits). Ligulate florets pink. Achenes up to 3.5 cm long, 
narrowed into beak, scabrous above, reddish on maturity; pappus dirty- 
66 yellow, wholly deciduous, pappus bristles connate at base, plumose, barbed 
above. Flowering June to August. 

Stony and clayey steppe slopes — Soviet Central Asia: Syr-Darya 
(Karatau), Tien Shan, Pamiro-Alai (including Shugnan). General 
distribution: Afghanistan. Described from cultivated specimens raised from 
seed in the Petersburg Botanical Garden. Type in Leningrad. 

Note. In the opinion of Popov, this wonderful species represents a 
hybrid between Tragopogon ruber and S. inconspicua{l). To me this 
opinion appears to be unfounded. The Kazakhstan specimens differ by 
having narrower Ught green or glaucous leaves, densely leafy stems, and 
a xerophilous appearance (5. tragopogonoides ssp. sclerophylla Krasch. 
and Lipsch.). 

28. S. petrovii Lipsch. in Lipsch. Sist. Zam ob Aziat. Skorts. (1932) 
20; Lipsch. in Fedde, Repert. XXXIII (1933) 167; Pavlov in Tr. po Prikl. 
Bot. Ser. 1, 2 (1937) 287; Lipsch. Fragm. Monogr. Scorzonera II (1939) 
79; Kovalevskaja in FI. Uzb. VI, 437.— Ic: Lipsch. op. cit. (1932) 35, 
Fig. 11; Pavlov in Zhum. "Sov. Kauchuk", 3 (1932) 27; Lipsch. op. cit. 
(1939) Plate 13. 

Perennial (? biennial). Plant 10-30 cm high; root collar covered with 
brown remnants of leaf sheaths. Stem terminating in more or less 
compressed candelabrumlike, corymbose inflorescence; branches of 



66 

inflorescence arcuate, densely leafy. Basal leaves herbaceous, narrowly 
linear, 0.2-0.4 cm wide, flat or partly incurved, glaucous-green, somewhat 
pubescent, with distinct, white, hyaline midrib beneath (width highly 
variable), somewhat arcuate, sometimes significantly exceeding stem (with 
inflorescence). Capitula solitary on all branches of corymbose 
inflorescence, 3.5-6.5 cm long, including ligulate florets. Involucre slightly 
arachnoid-pubescent, many-rowed, involucral bracts unequal; outer bracts 
deltoid-triangular, smaller; middle ones ovate-deltoid, somewhat narrowed 
above, broadly scarious along margin, two times as long as outer ones; 
inner ones linear-lanceolate, much longer than outermost. Ligulate florets 
pink. Achenes up to 3.5 cm long, glabrous, narrowed above into beak, 
with smooth or scabrous ribs, reddish when mature; pappus dirty yellow, 
deciduous; pappus bristles connate at base into a ring, plumose, glabrous 
at apex, barbed. Flowering June to July. 

Stony and gravelly mountain slopes. — Soviet Central Asia: Syr-Darya 
(Karatau Range), Tien Shan (western). Endemic. Described from Karatau. 
Type in Moscow. 

29. S. baldshuanica Lipsch. in ByuU. Mosk. Obshch. Isp. Prir. Otd. 

Biol. XLII, 2 (1933) 154; Lipsch. Fragm. Monogr. Scorzonera II (1939) 

80; Kovalevskaja in Fl. Uzb. VI, 437 (pro syn. S. tragopogonoides Rgl. 

67 and Schmalh.). — Ic: Lipsch. op. cit. (1933) 155, Fig. 1; Lipsch. op. cit. 

(1939) Plate 14. 

Perennial. Plant small, 5-9(13) cm high, glaucescent, pubescent plant. 
Root collar covered with brown sheaths of basal leaves. Stem solitary, 
slightly inclined, sulcate, usually dark red, partly arachnoid-pubescent, 
with three to five flowering shoots above. Leaves pubescent, glaberscent, 
flat or folded; basal leaves up to 15 cm long, herbaceous, linear, narrow, 
0.1-0.2(0.5) cm wide, at base somewhat broadened into sheaths; cauline 
leaves reduced, often undulate-crimped along margin. Capitula three to 
five, on short, usually tomentose peduncles, cylindrical, 3-3.5 cm long at 
flowering, up to 5 cm in early fruiting. Involucre many-rowed, arachnoid 
or white-tomentose; involucral bracts blackish-violet or with black spot, 
arachnoid pubescent, glabrescent; outer bracts smaller, deltoid, sometimes 
with herbaceous bent cusps; middle ones larger, ovate-deltoid, usually 
obtuse; inner ones longest, oblong-lanceolate. Ligulate florets pink. Mature 
achenes unknown. Flowering July. 

Stony and gravelly slopes of mountains. — Soviet Central Asia: Pamiro- 
Alai. Endemic. Described from Baldzhuan. Type in Leningrad. 

Section 5. Fibrillosae Nakai in Report. Inst. Sc. Res. Manch. I, 6 
(1937) 171. — Perennials. Root collar densely covered with dark brown 



67 

fibers of dead leaf petioles. Leaves crowded near stem base. Stems scape- 
like; with fewer (one to three) leaves. Florets yellow. Achenes smooth or 
pubescent. Pappus bristles plumose. 

Series 12. Subacaules Lipsch. in Addenda XXVIII, 720. — Plants more 
or less pubescent, almost stemless or scapelike, short, more or less 
pubescent. Capitula large, up to 5 cm long. Involucral bracts somewhat 
pubescent. Plant of alpine zone. 

30. S. subacaulis Lipsch. Byull. Mosk. Obshch. Isp. Prir. Otd. Biol. 
XLII, 2 (1933) 160; in Fedde, Repert. XXXIII (1933) 168; Fragm. 
Monogr. Scorzonera I (1935) 112. — S. austriaca var. intermedia Rgl. in 
Bull. Soc. Nat. Mosc. XL, 3 (1867) 170. — S. austriaca var. subacaulis 
Rgl. in Tr. Peterb. Bot. Sada, VI, 2 (1880) 323.— Ic: Lipsch. op. cit. 
(1933) 160, Fig. 6-7. 

Perennial. Root vertical, cylindrical; collar sparsely covered with 
blackish-brown hairs and sheaths of basal leaves, sheaths more or less 
broadly scarious along margin. Plants almost "stemless" or scape-like with 
solitary stems, less often two, (1)3.5-10(18) cm high, arachnoid-pubescent, 
especially below capitulum, few-leaved (one or two). Basal leaves linear, 
herbaceous, 2-4 mm wide, spreading or often vertical, usually longer 
68 than scape or as long, flat, with three indistinct veins (only midrib more 
or less prominent, broad), margin weakly scabrous, hyaline; cauline leaves 
one or two, membranous, scaly, sessile, lanceolate. Capitula solitary, large, 
broadly cyUndrical, 4(4.5)-5(5.5) cm long (including ligulate florets). Base 
of capitulum often enclosed by bract-like upper cauline leaf, similar in 
shape to outer involucral bracts. Involucre many-rowed, more or less 
arachnoid-pubescent, sometimes glabrescent; outer bracts deltoid or ovate, 
with broadly membranous margin; inner bracts oblong-lanceolate, usually 
obtuse. Ligulate florets yellow, with dark red parallel veins, pinkish when 
dry. Achenes slightly curved, 8-10 mm long, 1.5 mm wide, glabrous, 
striate, smooth; pappus dirty white, its bristles plumose, toothed-scabrous 
above. Flowering June to July. 

Alpine zone, in alpine meadows (Kobresian, etc.) and screes. — Soviet 
Central Asia'. Tien Shan. General distribution: Dzhungaria-Kashgaria. 
Described from Kegen District — Labazy Mountains in Sardzhas Valley. 
Type in Moscow. 

Note. It is very probable that S. subacaulis Lipsch. represents a 
polyploid alpine race whose progenitor was S. austriaca Willd. If this 
assumption is confirmed by cytological analysis, it would be more correct 
to include S. subacaulis Lipsch. under series Austriacae Lipsch., and 
eliminate the series Subacaules Lipsch. 



68 

Series 13. Austriacae Lipsch. in Addenda XXVIII, 720. — Plants 
glabrous, usually with stem. Involucral bracts glabrous. A plant of steppe, 
forest-steppe, and forest regions. 

31. S. crispa M.B. Fl. taur.-cauc. II (1808) 234; DC. Prodr. VII, 1. 
120; Ldb. Fl. Ross. II, 2, 794. — S. austriaca var. crispa (M.B.) Trautv. in 
Bull. Soc. Nat. Mosc. 39, 2 (1866) 382; Boiss. Fl. or. Ill, 770.— 5. 
prescottii Compere ex DC. Prodr. VII, 1 (1838) 120, nomen. — S. austriaca 
ssp. crispa (M.B.) Lipsch. and Krasch. in Lipsch. Fragm. Monogr. 
Scorzonera I (1935) 118.— Exs.: Gallier, Iter. Taur. tertium a. 1900, No. 
788. 

Perennial. Plant glabrescent; root collar with distinct hairs, often 
producing series of lateral leaf rosettes. Stems (2)10-20(28) cm high, 
glabrous, fistular, mostly straight, branched, with two to four capitula, 
less often simple and with single capitulum; lateral branches arising from 
axils of scaly cauline leaves. Leaves glaucous, coriaceous, mostly ovate- 
lanceolate, or oblong-lanceolate, up to 5 cm wide (usually 1.5-3 cm), 
sometimes falcate, narrowed toward both ends (apex attenuate into hood), 
with somewhat distinct veins, along margin with cartilaginous and 
somewhat toothed stripe, undulate-crimped; basal leaves mostly long- 
petiolate, petioles basally broadened, sometimes floccose on inner side 
when young, later becoming glabrous; capitula 2.5-3 cm long (at 
69 flowering), often covered with scaly bracts. Involucre glabrous; bracts 
membranous along margin, mostly subobtuse; outer bracts reduced. 
Ligulate florets yellow, exceeding involucre. Outer achenes somewhat 
curved, 9-10 mm long, glabrous (sometimes with crown of hairs only at 
tip), ribbed, sometimes with barely visible tubercles along ribs, usually 
without them. Flowering April to May. (Plate IV, Fig. 2.) 

Limestone hills, slopes, rocks. — European Part: Crimea; Soviet 
Central Asia: Balkhash Region (Ulutau Range, hilly terrain of Kazakhstan). 
Endemic. Described from Crimea. Type in Leningrad. 

Note. I have included specimens from the Ulutau Range and the 
adjacent outlier mountains of Kazakhstan under this species. They also 
are characterized by glaucescent plants, oval, broadly lanceolate or oblong- 
lanceolate leaves with an aristate tip, and are identical to the Crimean 
plants. In the herbarium of the Botanical Institute, Academy of Sciences 
of the USSR, I had earlier identified the Kazakhstan specimens of this 
species as S. austriaca var. ulutauensis Lipsch. (ined.). The abundant fresh 
material has convinced me that the Crimean and Ulutau specimens are 
the same species. 

32. S. ruprechtiana Lipsch. and Krasch. in Lipsch. Fragm. Monogr. 
Scorzonera I (1935) 121, pro syn. — 5. (austriaca) glabra Rupr. Flor. 



69 

Samojed. cisuralens. (1845) 11, in textu; Kiyl. Fl. Zap. Sib. XI, 2984.— 
S. austriaca var. glabra Rupr. in Fl. Sev. Urala (1856) 12 and 40. — S. 
austriaca subsp. glabra (Rupr.) Lipsch. and Krasch. in Lipsch. op. cit. 
(1935) 121.— Ic: Lipsch. op. cit. (1935) Plate 44. 

Perennial. Root cylindrical, long. Root collar densely hairy. Stem 
(15)25-50 cm high and 3-6 mm thick, sulcate, hollow, glabrous, often 
more or less swollen below capitulum. Basal leaves broadly lanceolate to 
oblong-ovate, narrowed into winged petiole, together 20-30(40) cm long 
and (0.9) 1.2-3.5(5) cm wide, glabrous, along margins flat; cauline leaves 
two or three, broadened at base; lower leaves broadly lanceolate; upper 
leaves small, scaly. Capitula solitary, large, (3)3.5^(4.5) cm long with 
florets; involucre 1.2-1.5 cm wide; outer bracts broadly ovate, glabrous; 
inner bracts oblong. Ligulate florets yellow. Achenes 10-13(15) nmi long, 
somewhat curved, glabrous, smooth, ribbed, with ring of hairs at apex; 
pappus dirty white. Flowering June to July. 

Mountain peaks, slopes and rocks, limestone deposits; gypsum, sandy 
scarps. — European Part: Dvina-Pechora, Volga-Kama, Volga-Don 
(Zhiguli); Western Siberia: Upper Tobol (alpine zone of Urals up to GubeU 
Mountains); Eastern Siberia: Angara-Sayans, Dauria. Described from 
northern Urals. Type in Leningrad. 

33. S. pratorum (Krasch.) Stankov in Stank, and Tal. Opred. Vyssh. 
Rast. (1949) 687. — S. austriaca var. pratorum Krasch. in Lipsch. Fragm. 
Monogr. Scorzonera I (1935) 116 and in H. Yugo-Vost. VI (1936) 451.— 
5. austriaca ssp. pratorum (Krasch.) Lipsch. in sched. — Ic: Lipsch. op. 
cit. (1935) Plate 42. 

70 Perennial. Plant glaucous-green. Stems tall, 25-40(50) cm high, 

usually dichotomously branched above. Leaves oblong-lanceolate, (1)1.5- 
2.5(4) cm wide; basal leaves long-petiolate; cauUne leaves sessile. Capitula 
large, (2)2.5 cm (at flowering) and 3-3.5 cm (at fruiting) long; achenes 
9-1 1 mm long, glabrous, smooth, ribbed. Flowering June to July. 

Saline meadows, deltas. — European Part: Trans- Volga, Lower Volga; 
Western Siberia: Upper Tobol (lower part), Irtysh. Type in Leningrad. 

34. S. austriaca Willd. Sp. pi. Ill, 3 (1803) 1498; Ldb. Fl. Ross. II, 
2, 792; Boiss. Fl. or. Ill, 770; Schmalh. Fl. II, 143; Lipsch. Fragm. 
Monogr. Scorzonera I, 107; Krasch. in Fl. Yugo-Vost. VI, 450; Kryl. Fl. 
Zap. Sib. XI, 2981; Lipsch. in Spisok Rast. Gerb. H. SSSR XIII, 130.— 
S. humilis auct. non L.; Jacq. Enum. stirp. Vindob. (1762) 139 and R. 
Austr. I (1773) t. 36; DC. Prodr. VII, 1 (1838) 120 p. p.— 5. graminifolia 
auct. non L.: Ldb. Fl. Alt. IV (1833) 161.— Ic: Rchb. Ic. R. Germ. XIX, 
t. 1383; Javorka and Csapody, iconogr. fl. Hungar. No. 3996. — Exs.: Fl. 
exs. Austro-Hung, No. 2243; GRF No. 3996. 



70 

Perennial. Whole plant glabrous. Root cylindrical, vertical, dark 
brown. Root collar covered with numerous dark brown fibers of dead 
petioles. Stems solitary or a few, (5)10-30 cm high, simple or branched 
above, sparsely leafy (usually bearing one to three leaves). Leaves always 
variable in width, linear, linear-lanceolate or broadly lanceolate, flat along 
margin or more or less undulate-crimped, with several veins; basal leaves 
narrowed into petioles; cauline leaves one to three, reduced, linear or 
Unear-lanceolate, sessile, with somewhat broadened base. Capitula solitary 
or two to four, broadly cylindrical, 1.8-2.5 cm long and 0.5-1(1.5) cm 
wide; involucre glabrous; outer bracts ovate or deltoid-ovate, obtuse 
or acute; innear lanceolate, obtuse or subobtuse with scarious margin. 
Ligulate florets yellow, one and one-half to two times as long as involucre. 
Achenes glabrous, with ring of hairs only below pappus, ribbed, ribs 
smooth or tuberculate; pappus dirty white, as long as achene. Flowering 
June to July. 

Steppes, stony-gravelly slopes of hills and mountains. — European 
Part: Bessarabia, Middle Dnieper, Black Sea Region, Lower Don, Lower 
Volga, Trans- Volga, Volga-Kama; Western Siberia: Upper Tobol, Irtysh, 
Altai; Eastern Siberia: Angara-Sayans, Lena-Kolyma; Soviet Central Asia: 
Aralo-Caspian, Balkhash region. General distribution: Central Europe, 
Mediterranean, Balkans-Asia Minor, Mongolia. Described from Austria. 
Type in Berlin. 

Note. A highly polymorphic species, varying in shape and width of 
leaves, growth, size and number of capitula, achene sculpture, etc. On 
the basis of working through all the material from the USSR, except for 
a series of geographic (5. crispa M.B., S. ruprechtiana Lipsch. and 
Krasch., S. dianthoides (Lipsch. and Krasch.) Lipsch., S. curvata (Popl.) 
72 Lipsch.) and ecological (5. pratorum (Krasch.) Stank.) races examined 
separately, the following key to the varieties of 5". austriaca s. 1. can be 
proposed with the caution that a large number of transitional forms is 
found, which do not fit into my proposed classification of this species. 

1. Mature achenes densely covered with tubercles along ribs 

var. verrucosa Lipsch. and Krasch. 

-I- Achenes smooth, without tubercles, or outer achenes slightly 
tuberculate below along ribs 2. 

2. Capitula two to four on single stem. Peduncles thick, straight 

var. triflora (Fisch.) Lipsch. and Krasch. 

-I- Capitula solitary 3. 

3. Leaves hnear, narrow (1-3.5 mm wide) 

var. tenuifolia Lipsch. and Krasch. 

+ Leaves broader 4. 

4. Leaves glaucous, hard, cartilaginous, 1.5-3 cm wide, with apical 



71 




71 Plate VI. 

1 — Scorzonera armeniaca (Boiss. and Huet) Boiss.; 2 — 5. biebersteinii Lipsch. 



72 

spinule, highly undulate-crimped along margin 

; S. crispa M.B. (see above). 

+ Leaves less hard, curved, broadly lanceolate, less often linear, 0.4- 
1 .5 cm wide, partly folded, along margins slightly undulate-crimped 
var. altaica Serg. 

A detailed description of these varieties can be found in the cited 
publications of Lipschitz (1935) and Krylov (1949). 

A large number of transitional specimens is known from the 
Transbaikal Region and Yakutia, which cannot be accommodated in the 
proposed classification scheme of 5. austriaca. These forms probably are 
of hybrid origin, involving S. radiata Fisch. Additional collections and 
observations are needed. 

35. S. dianthoides (Lipsch. and Krasch.) Lipsch. comb. nova. — 5. 
austriaca var. dianthoides Lipsch. and Krasch. in Lipsch. Fragm. Monogr. 
Scorzonera I (1935) 116.— Ic: Lipsch. op. cit. (1935) Table 41. 

Perennial. Stems 20-25 cm high, slender, branched above; flowering 
branches two to four, slender, bent. Leaves Unear, 2-4 mm wide, or linear- 
lanceolate, 7-12 mm wide. Capitula narrowly-cylindrical, 1.5-2 cm long, 
3-5 mm wide. Ligulate florets yellow, pinkish when dry. Achenes 7-8 
mm long, ribbed, smooth; pappus white, plumose, its bristles barbed above. 
Flowering May. 

Crevices in rocks. — Western Siberia: Irtysh (Arkadian Mountains in 
Semipalatinsk Region). Endemic. Described from the above place. Type 
in Leningrad. 

36. S. curvata (Popl.) Lipsch., comb. nova. — S. austriaca var. curvata 
Popl. in Tr. Bot. Muz. Akad. Nauk XV (1916) 38; Lipsch. 

73 Fragm. Monogr. Scorzonera I (1935) 114. — S. humilis angustifolia Turcz. 
in DC. Prodr. VII, 1 (1838) 120 and in Herb. Acad. Sc. URSS.— 5. humilis 
a. linearifolia DC. op. cit. (1838) 120 (excl. syn. S. angustifolia Thom. 
and S. graminifolia a. in Ldb. Fl. Alt. IV (1833) 161). — S. austriaca p. 
linearifolia DC. ex Turcz. in Bull. Soc. Nat. Mosc. XXI, 3 (1848) 95 (Fl. 
baic.-dahur.). — S. austriaca var. pusilla Turcz. nomen in herb, and ex 
Lipsch. op. cit. (1935) 114. — Ic: Poplavskaja, op. cit. (1916) page 39, 
Fig. 1. 

Perennial. Small plant, 2-6(10) cm high. Leaves fihform, narrow (1- 
1.5 mm wide), glaucous, usually helical, longer than stems (together with 
capitulum). Compared with small size of the whole plant, capitula fairly 
large, (2)2.5-3 cm long. Involucre glabrous; outer bracts deltoid-lanceolate, 
acuminate, inner oblong-lanceolate, usually somewhat obtuse; all bracts 
smaller and narrower than typical S. austriaca. Ligulate florets yellow. 
Flowering May to June. 



73 

Steppes. — Eastern Siberia: Dauria, Angara-Sayans. General 
distribution: Mongolia. Described from Dauria. Type in Leningrad. 

Note. The specimens from the Angara-Sayan District differ from the 
Daurian ones: they are transitional to S. austriaca var. tenuifolia Lipsch. 
and Krasch. According to the practice followed in the Flora of the USSR, 
I am giving this taxon a binomial name. 

Section 6. Pulvinares (Boiss.) Lipsch. in Lipsch. Fragm. Monogr. 
Scorzonera I (1935) 24. — Subsection Pulvinares Boiss. Fl. or. Ill (1875) 
756. — Plants usually small, forming dense tussocks. Stems scapelike, 
leafless or few-leaved. Leaves linear or lanceolate. Pappus bristles stiff, 
fragile, often colored, plumose, barbed-scabrous above. 

Series 14. Filifoliae Lipsch. in Addenda XXVIII, 720.— Stems 
scapelike, few. Leaves narrow, filiform-linear. Root collars sparsely 
covered with sheaths. Achenes glabrous or pubescent. 

37. S. filifolia Boiss. Fl. or. Ill (1875) 774; Medved. in Tr. Tifl. Bot. 
Sada, XVIII, 2 (1918) 306; Grossh. in bot. Shorn. Azerb. GNU, I (1932) 
54; Grossh. Fl. Kavk. IV (1934) 240; Lipsch. Fragm. Monogr. Scorzonera 
I (1935) 28, p. max. p.; II (1939) 14-15; Sosnowsky in Zam. po Sist. i 
Geogr. Rast. Tbil. 13 (1947) 62-64, p. p.; Grossh. Opred. Rast. Kavk. 
510; Papava in Fl. Gruzii, VIII, 651; Isaev in Fl. Azerb. VII, 522.-5. 
filifolia var. vegetiorTrsLUtv. in Tr. Peterb. Bot. Sada, X (1887) 119.— 5. 
rubriseta Lipsky in Zap. Kiev. Obshch. Estestv. XI (1891) 52; in Tr. 
Peterb. Bot. Sada, XIII, 315, p. max. p.; Grossh. op. cit. (1934) 241; 
Lipsch. op. cit. I (1935) 30; II (1939) 14, p. p. — S. filifolia var. rubriseta 
(Lipsky) Grossh. op. cit. (1932) 55. — S. charadzae Papava in Fl. Gruzii, 
VIII (1952) 651 (descr. georg.) and t. 435.— Ic, Lipsch. op. cit. (1935) 
Plate 8.— Exs.: Herb. fl. Cauc, No. 396, p. p. 
74 Perennial. Plant with many-headed rhizome, forming dense tussocks. 

Root collar covered with sheaths; on inner side covered with woolly 
growth. Scapes ascending, 7.5-10(25) cm long (including capitula), 
slender, sulcate, slightly pubescent (glabrescent), with one or three or 
four capitula, and a few (one to three) scaly leaves; almost all leaves 
basal, numerous, filiform-linear, rather narrow, 0.5-1 mm wide, weakly 
pubescent, or almost glabrous. Capitula turbinate, 1.1-1.6 cm long; 
involucre weakly pubescent; outer involucral bracts small, lanceolate; inner 
ones lanceolate, oblong, several times as long as outer; all bracts with 
narrow red stripe along margin. Ligulate florets yellow, exceeding 
involucre by one-third length. Achenes glabrous, narrowly cylindrical, 
with thin ribs; pappus dirty white or reddish-brown; pappus bristles stiff, 
fragile, plumose, barbed-scabrous above. Flowering June to August. 



74 

Stony and limestone slopes. — Caucasus: Dagestan, Eastern 
Transcaucasia. Endemic, described from Dagestan. Type in Leningrad. 

Note. The varieties being recognized are: var. incanescens Sosn. op. 
cit. (1947) 63, distinguished by tomentose-lanate involucral bracts, 
peduncles, and leaves; var. vegetior Trautv. (= S. rubriseta Lipsky, ex 
parte) — up to 25 cm high, often with branched stems and three or four 
capitula, pappus usually reddish-brown. I did not see the original specimens 
from which S. charadzae Papava was described (they are preserved in 
Tbilisi). The species diagnosis is given in the Georgian language and is 
not accessible to me. Judging from the figure of S. charadzae, it represents, 
in my opinion a large variety of S. filifolia and does not deserve to be 
treated as an independent species. S. filifolia is a highly variable species 
in stem height, nature of branching, leaf width, pappus color, etc. 

38. S. lipskyi Lipsch. Fragm. Monbgr. Scorzonera II (1939) 15. — 5. 
filifolia auct. non Boiss.; Lipsch. op. cit. (1935) 28, p. min. p. — S. filifolia 
var. lipskyi (Lipsch.) Sosn. in Zam. po Sist. i Geogr. Rast. Tbil. 13 (1947) 
63; Grossh. Opred. Rast. Kavk. 510. 

Perennial. Very similar to S. filifolia Boiss. Stem scapelike, up to 25 
cm high, sparsely leafy (cauline leaves one to three); leaves linear- 
lanceolate, very narrow (up to 1 mm wide). Capitula turbinate, 1.2-1.5 cm 
long, pubescent; pappus usually reddish-brown. Flowering June to July. 

Stony and limestone slopes. — Caucasus: Dagestan. Endemic. • 

Note. Contrary to the opinions of Sosnowsky and Grossheim, for the 
present I maintain the separate status of this species, which is quite close 
to S. filifolia Boiss., but differs from the latter and other members of 
section Pulvinares by the presence of pubescence over the entire length 
of the achene. Although both species were described from the same area 
(Dagestan), only careful observations of both taxa in their native habitats, 
on more extensive material than that in my hands, will make it possible 
to resolve finally the "enigma" concerning the separateness of these close 
species. 

75 39. S. seidlitzii Boiss. Fl. or. Ill (1875) 775; Seidlitz in Tr. Bot. 

Sada, Yur'ev Univ. IV (1903) 66; Medved. in Tr. Tifl. Bot. Sada, XVIII, 
2 (1918) 306; Grossh. in Bot. Shorn. Azerb. GNU, I (1932) 54; Grossh. 
Fl. Kavk. IV (1934) 240; Lipsch. Fragm. Monogr. Scorzonera I (1935) 
26; Grossh. Opred. Rast. Kavk. 510; Papava in Fl. Gruzii, VIII, 648; Juz. 
in Spisok Rast. Gerb. Fl. SSSR XIV, 77; Dmitrieva, Opred. Rast. Adzhar. 
(1960) 287.— Ic: Lipsch. op, cit. (1935) Plates 6, 7; Dmitrieva, op. cit. 
(1960) Plate 34.— Exs.: GRF No. 4100. 

Perennial. Rhizome vertical, with a few caudices, forming tussocks. 
Root collar densely covered with lanate sheaths of basal leaves. Basal 
leaves linear, flat or grooved, 1 .5—2.5 mm wide, slightly attenuate toward 



75 

apex, densely villous, sinuate, hairs divergent, white, becoming yellow in 
herbarium. Scapes ascending from base (4)8-18(20) cm long (including 
capitula), with single capitulum (less often bifurcated in lower part and 
then with one or two branches), sulcate, pubescence similar to that of 
leaves, but hairs usually longer. Capitula turbinate, 1.3-2(2.4) cm long, 
including ligulate florets; involucre woolly-pubescent; outer bracts 
particularly densely hairy, foUaceous with uncinate apices; inner bracts 
blackish-brown, with carinate midrib, somewhat coriaceous, linear- 
lanceolate, two times as long as outer ones. Ligulate florets lemon-yellow, 
pinkish when dry, one and one-half times as long as outer involucral 
bracts. Achenes glabrous, 5-6 mm long, slightly curved, cylindrical, 
reddish-brown when mature, with smooth longitudinal ribs; pappus dirty 
yellow or often reddish-brown, as long as achene, its bristles stiff, fragile, 
plumose, barbed-scabrous above. Flowering June to August. (Plate V, 
Fig. 1) 

Subalpine meadows, stony gravelly slopes. — Caucasus: Eastern 
Transcaucasia, Southern Transcaucasia. General distribution: Armenia and 
Kurdistan. Described from Alagyoz. Type in Geneva. 

Note. It differs from the related species S. filifolia Boiss. by having 
wider leaves, longer, flexuous, divergent hairs scattered over the whole 
plant, a woolly pubescent involucre, and longer involucral bracts. 

Series 15. Rigidae Lipsch. in Addenda XXVIII, 720.— Stems 
scapelike, taller, numerous, slender, and erect. Root collar densely covered 
with rigid sheaths. Leaves narrow, linear. 

40. S. rigida Auch. ex DC. Prodr. VII, 1 (1838) 123; Boiss. Fl. or. 
Ill, 774; Post, Fl. Syr., Palest, a. Syn. (1896) 494; Grossh. in Bot. Sbom. 
Azerb. GNU, I (1932) 54; Grossh. Fl. Kavk. IV (1934) 240; Lipsch. 
Fragm. Monogr. Scorzonera I (1935) 30; Lipsch. in Spisok Rast. Gerb. 
SSSR, XIV, 129; Isaev in H. Azerb. VIII, 522.— Ic: Lipsch. op. cit. 
(1935) Plates 9, 10; Figs. 1, 2.— Exs.: GRF No. 4200. 
76 Perennial. Plant with vertical cylindrical rhizome, branched at apex, 

producing several caudices crowded together; collar of caudices very 
densely covered with hard leaf sheaths, lanate on inner side. Stems (20)30- 
40 cm high, numerous, slender, hard, erect, spreading, simple or 
dichotomously branched, glaucescent, sericeous; later almost glabrous, 
weakly leafy, much longer than basal leaves. Leaves narrow, filiform- 
linear, glaucescent-green, flat or incurved, flexuous, pubescent; basal leaves 
broadened at base into stramineous, glossy sheaths, lanate on inner side; 
cauline leaves few, reduced, filiform, sessile, at base slighdy broadened. 
Capitula small, 1-1.5 cm long, turbinate, solitary, terminal on stem and 
branches. Involucre pubescent, bracts weakly carinate, slightly 
membranous along margin; outer bracts very small, deltoid, acuminate; 



76 

inner bracts linear-lanceolate, subobtuse, three to four times as .long 
as outer. Ligulate florets lemon-yellow, longer than involucre by one- 
third. Achenes 6-8 mm long, glabrous, with obtuse ribs; pappus dirty 
white, its bristles fragile, plumose, barbed-scabrous above. Flowering July 
to August. 

Stony mountain slopes. — Caucasus: Southern Transcaucasia. General 
distribution: Armenia and Kurdistan. Described from West Asia. Type in 
Geneva, isotype in Leningrad. 

Series 16. Koslovskyianae Lipsch. in Addenda XXVIII, 720. — Stems 
scapelike, leafy. Leaves lanceolate. Pappus hairs reddish-brown. 

41. S. koslovskyi Sosn. in Lipsch. Fragm. Monogr. Scorzonera I 
(1935) 42 (descript. lat.); Grossh. Fl. Kavk. IV (1934) 241 (descript. 
ross.).— 5. debilis Sosn. in Tr. Tbil. Bot. Inst. II (1938) 222; Grossh. 
Opred. Rast. Kavk. 510; Papava in Fl. Gruzii VIII, 648. — Ic: Lipsch. op. 
cit. (1935) Plate 15. 

Perennial. Plant with thick, vertical, dark brown rhizome, with a few 
caudices and indurescent remnants of stem bases and basal leaves in upper 
part. Stems solitary or up to five, 17-20 cm high, slightly bent, usually 
dichotomously branched above, and then with two capitula, distinctly 
angular, sulcate, more or less leafy, white tomentose, especially in lower 
part, glabrous or with scattered hairs above. Leaves hard, lanceolate, more 
or less pubescent or almost glabrous, with three to five veins; basal leaves 
lanceolate, 4-7 mm wide and 12-15 cm long, narrowed into sulcate, 
tomentose petiole; acuminate, with callous cusp; cauline leaves up to 10, 
lanceolate, reduced, lowermost narrowed into short petiole, higher ones 
almost sessile; all leaves slightly falcate. Capitula solitary terminal 
77 on stems or dichotomous, flexuous, axillary branches, cylindrical or 
turbinate, 1.8-2 cm (at flowering), and up to 2.5 cm long (at fruiting). 
Involucre pubescent, involucral bracts often reddish, narrowly membranous 
along margin; outer bracts small, lanceolate, acuminate, often with deflexed 
cusps; inner bracts oblong-lanceolate, deltoid, two to two and one-half 
times as long as outer ones. Ligulate florets yellow, with reddish veins 
when dry, longer than involucre. Achenes narrowly cylindrical; 7-8 mm 
long, stramineous, glabrous, with obtuse longitudinal ribs; papus reddish- 
brown, slightly longer than achenes; its bristles fragile, plumose, barbed- 
scabrous above. Flowering July to August. 

Stony-gravelly substrates. — Cacucasus: Eastern Transcaucasia 
(Akhalkalaki Plateau, Tetrob Range). Endemic. Described from the above 
place. Type in Moscow; isotype in Tbilisi. 



77 

Note. On the basis of the leafy stem and the pubescence, this species 
can be formally included under section Tomentosae (Boiss). Lipsch. 
However, the combination of other characters (nature of the achene, 
pappus, involucre, and general appearance) compels me to include 5. 
koslovskyi Sosn. in section Pulvinares, to the members of which, in my 
opinion, it is more closely related. It is a relict endemic plant of the 
Akhalakalaki Region of the Georgian SSR, where it is found scattered as 
isolated plants. 

Section 1. Scorzonera. — Radiatae Nakai in Report. Inst. Sc. Res. 
Manch. I, 6 (1937) 169. — Root perennial [sic; many-headed], with scales 
and not hairs near collar. Leaves basal and cauline. Stems one-headed. 
Outer involucral bracts reduced or all almost equal. Achenes smooth. 
Pappus bristles plumose. 

42. S. radiata Fisch. in Ldb. Fl. Alt. IV (1833) 160; DC. Prodr. 
VIII, 1, 122; Ldb. Fl. Ross. II, 2, 793; Turcz. in Bull. Soc. Nat. Mosc. 
XXI, 3 (1848) 96 (Fl. baic.-dahur.); Kryl. Fl. Alt. 3, 733; Kom. and Alls. 
Opred. Rast. Dalnevost. Kr. II, 1092; Lipsch. Fragm. Monogr. Scorzonera 
I (1935) 124; Kryl. Fl. Zap. Sib. XI, 2984; Lipsch. in Spisok Rast. Gerb. 
Fl. SSSR, XIII, 131.— 5. radians Fisch. ex Bess, in Flora XVII (1834) 1 
Beibl. 18.— Exs.: GRF Nos. 1225, 3999. 

Perennial. Root cylindrical, vertical or ascending, one- or many- 
headed, in latter case with rosette of basal leaves and a few (two to six) 
simple stems, each terminating in capitulum. Root collar covered with 
brown membranous sheaths (or their remnants), never fibrous. Stems (3)5- 
40(50) cm high, straight, weakly sulcate, more or less arachnoid-tomentose, 
sometimes floccose-pubescent, especially at base and apex, often almost 
glabrous in adult specimens, widi a few (one to three) leaves. Basal leaves 
linear or linear-lanceolate; (3.5)5-30 cm long and 3-15(20) mm wide, 
acuminate, flat, with three to five veins, narrowed toward base into petiole, 
glabrous or slightly arachnoid-pubescent; cauline leaves (one to three) 
reduced, (1.5)3-8(12) cm long, sessile, linear, or lanceolate; uppermost 
78 leaves often scaly. Capitulum large, soUtary, 2.5-3.5(4) cm long (including 
ligulate florets). Involucre glabrous or slightly arachnoid-pubescent, 
imbricate; outer bracts ovate-lanceolate, smaller; inner bracts linear-oblong, 
longer; all bracts slightly membranous along margins, obtuse and with 
red spot at apex. Ligulate florets light yellow, two to two and one-half 
times as long as involucre. Achenes 10-12 mm long, glabrous, smooth, 
ribbed; pappus dirty white; its bristles plumose, five of them longer and 
barbed above. Flowering May to August (Plate V, Fig. 2). 



78 

Alpine and forest zones of mountains, less often in the plains. Stony- 
gavelly and moss-lichen tundras, alpine and forest meadows, meadow- 
covered slopes, edges of coniferous forests. — Western Siberia: Irtysh 
(eastern part), Altai; Eastern Siberia: Yenisei, Lena-Kolyma, Angara- 
Sayans, Dauria; Far East: Anadyr, Okhotsk, Zeya-Bureya, Uda area, 
Ussuri, Sakhalin. General distribution: Mongoha, China. Described from 
Altai. Type in Leningrad. 

Note. Known from the Okhotsk Region is var. subacaulis Lipsch. 
and Krasch. (Lipsch. Fragm. Monogr. Scorzonera I (1935) 125) — a dwarf 
plant, almost stemless, 1-2 cm high, stems three, densely tomentose; leaves 
1-3 mm wide, slightly spreading, weakly uncinate at tips, distinctly longer 
than stems. From the alpine zone of the Sikhote Alin Range (Far East) is 
a plant in the Herbarium of the Botanical Institute, Academy of Sciences 
of the USSR labeled var. alpina Soczara — a shorter plant (10-15 cm 
high), with narrow leaves (0.4-0.8 cm wide). 

43. S. humilis L. Sp. pi. (1753) 790; Ldb. Fl. Ross. II, 2, 793; 
Schmalh. Fl. II, 143; Syreistsch. Illyustr. Fl. Mosk. Gub. Ill, 327; Hegi, 
111. Fl. VI, 2, 1057. —5. nervosa Gilib. Fl. Lithuan. Ill (1781) 230, non 
Trev.; Pers. Synops. II (1807) 360. — S. plantaginea Schleich. ex Gaud. 
H. helv. V (1829) 20; DC. Prodr. VII, 1, 119.— Ic: Rchb. Icon. Fl. Germ. 
XIX, t. 1383; Syreistsch. op. cit. 327; Hegi, op. cit. t. 276, fig. 4, 4a, fig. 
741, g-k; 742-744.— Exs.: Herb. Fl. Ingr. No. 356; Herb. Fl. Reipubl. 
Sov. Ucr. (1925) No. 100; Eesti Taimed. Tartu UlikooU Hot. muus. No. 
137. 

Perennial. Plant lanate, glabrescent. Root cylindrical; root collar 
covered with membranous scales. Stems solitary or three, (5)15-25(50) 
cm high, hollow, sulcate, simple or less often branched at apex, with a 
few (one to three) cauline leaves. Basal leaves long-petiolate, variable in 
shape and width; oblong-ovate or elliptical, less often lanceolate, with 
quite prominent veins, acuminate, entire; cauline leaves (one to three) 
sessile, lanceolate, or lanceolate-linear. Capitula large, 3.5-4(5) cm long 
(including ligules); peduncles usually lanate below capitulum. Involucre 
pubescent; outer bracts ovate-deltoid; inner ones longer than outer, 
lanceolate or oblong-lanceolate, obtuse and often with a red spot at apex. 
79 Ligulate florets light yellow, almost two times as long as involucre. 
Achenes 7-1 1 mm long, smooth, ribbed; pappus dirty-white, its bristles 
weakly plumose, longer ones barbed above. Flowering May to June. 

Meadows; among shrubs. — European Part: Baltic Region, Ladoga- 
Ilmen, Upper Dnieper, Upper Volga (southern part), Volga-Don, ?Middle 
Dnieper. General distribution: Scandinavia, Central Europe, Atlantic 
Europe. Described from Europe. Type in London. 



79 

Section 8. Parviflorae Lipsch. in Lipsch. Fragm. Monogr. Scorzonera 
II (1959) 112, pro subsect. — Plant of saline habitats. Leaves green or 
glaucous, more or less fleshy; stems without distinct virgate branches. 
Capitula small, with small flowers; ligulate florets slightly exserted from 
involucre; achenes glabrous, ribbed, smooth; pappus snow-white, pappus 
bristles plumose. 

Series 17. Parviflorae Lipsch. in Addenda XXVIII, 720. — Root collar 
not thickened. Stems simple, with one to three scaly leaves and one (less 
often two) terminal capitulum. 

44. S. parviflora Jacq. Fl. Ausr. IV (1776) 3; Willd. Sp. pi. Ill, 3, 
1500; Ldb. Fl. alt. IV, 161; DC. Prodr. VII, 1, 121; Ldb. Fl. Ross. II, 2, 
795; Boiss. Fl. or. Ill, 770; Maxim, in Bull. Acad. Sc. Petersb. XXXII, 4, 
(1888) 491; Schmalh. Fl. II, 143; Kryl. Fl. Alt. Ill, 734; Hegi, 111. Fl. VI, 
2, 1060; Grossh. Fl. Kavk. IV (1934) 242; Lipsch. Fragm. Monogr. 
Scorzonera II, 136; Kryl. Fl. Zap. Sib. XI, 2989; Lipsch. in Spisok Rast. 
Gerb. Fl. SSSR, XIII, 131; Isaev in Fl. Azerb. VIII, 523; Lipsch. in Fl. 
Turkm. VII, 307; Kovalevskaja in Fl. Uzb. VI, 438.-5. caricifolia Pall. 
Reise III (1776) 539 and Anh. 767; Willd. Sp. pi. Ill, 3, 1500; Claus in 
Gobel, Reise II, 184.— 5. halophila Fisch. and Mey. in DC. Prodr. VII, 1 
(1838) 122; Boiss. and Buhse in Nouv. Mem. Soc. Nat. Mosc. XII, 139.— 
S. schanginiana Fisch. ex Lipsch. Fragm. Monogr. Scorzonera II (1939) 
137, nomen.— Ic: Jacq. op. cit. t. 305; Pall. Reise III, 2 (1776) t. I/, fig. 
1; Pall. Putesh. Ill, 2 (1778) Plate I, i, fig. 3; Hegi, op. cit. fig. 745.— 
Exs.: Lang and Szovits, Herb. ruth. cent. II, No. 139; GRF No. 3998. 

Perennial or biennial. Plant wholly glabrous; root branched, many- 
headed, often producing rosette of radical leaves and one or several stems; 
root collar bare or covered with scales. Stems (10)15-^0 cm long, erect 
or ascending at base, usually hollow, simple, with single capitulum or 
less often weakly branched, and then with two or three capitula, sparsely 
leafy, with one to three scaly leaves. Leaves fleshy, slightly curved, with 
five to seven veins; basal leaves lanceolate or less often elliptical- 
lanceolate, 7-20 cm long, 5-15 nrmi wide, acuminate, narrowed at base 
into petiole broadened at very base into sheath; cauline leaves small, linear- 
lanceolate. Capitula solitary, less often two or three, rather small, 13-20 
mm long and 3-7 mm wide, broadened at fruiting. Involucre glabrous; 
outer bracts ovate-deltoid, acuminate; inner ones oblong-lanceolate, 
80 subobtuse, sometimes with blackish apical spot; all bracts narrow and 
scarious along margin. Ligulate florets yellow-waxy, very slightly longer 
than involucre. Achenes 7-9 mm long, glabrous, ribbed, ribs smooth; 
pappus snow-white, two times as long as achenes, plumose; five of its 
bristles longer, barbed above. Flowering May to July. 



80 

Solonchaks, saline meadows, seaside meadows, saline marshes. — 
European Part: Volga-Kama (southern part). Middle Dnieper, 
Volga-Don, Trans- Volga, ?Upper Dniester, Bessarabia, Black Sea Region, 
Crimea, Lower Don, Lower Volga; Caucasus: Ciscaucasia, Eastern and 
Southern Transcaucasia; Western Siberia: Upper Tobol, Irtysh; Soviet 
Central Asia: Aralo-Caspian, Balkhash Region, ?Kyzyl-Kum, Kara-Kum, 
mountainous Turkmenia, Amu-Darya, Syr-Darya, Pamiro-Alai, Tien Shan. 
General distribution: central Europe, eastern Mediterranean (Cyprus), 
Armenia and Kurdistan, Iran, Dzhungaria-Kashgaria, Mongolia. Described 
from Austria. Type in Vienna. 

Note. S. carcifolia is described from specimens collected from the 
Naryn sands and in the vicinity of Iletskaya Zashchita. Type in Leningrad. 
5". halophila is described on the basis of specimens from Iran. Type 
preserved there. 

Series 18. Mongolicae Lipsch. in Addenda XXVIII, 720. — Root collar 
thickened from numerous remnants of indurescent sheaths of basal leaves. 
Stems branched, leafy, with several (up to six) capitula. 

45. S. mongolica Maxim, in Bull. Acad. Sc. Petersb. XXXII, 4 (1888) 
492; Lipsch. Fragm. Monogr Scorzonera II (1939) 132. — S. mongolica 
var. putjatae Winkl. in Tr. Peterb. Bot. Sada, XIV (1898) 128.— .?5. 
fengtiensis Nakai in Report. Inst. Sc. Res. Manch. I, 6 (1937) 167. — Ic: 
Lipsch. op. cit. (1939) Plate 21. 

Perennial. Glaucous, glabrous plant. Root vertical, cylindrical; root 
collar sometimes two- or three-headed, covered with brown or stramineous 
indurescent sheaths of basal leaves. Stems more or less numerous, (2.5) 
6-20(40) cm high, branched above, straight, ascending at base or 
sometimes prostrate, glabrous, sulcate, leafy. Leaves fleshy, glaucous, 
rugose in herbarium, mostly falcate, with three to five indistinct veins; 
basal leaves lanceolate or linear-lanceolate, acuminate (cusp cartilaginous), 
narrowed into petiole broadened at base into sheath, floccose within; 
cauline leaves sessile, reduced in upper part of stem, linear-lanceolate, 
alternate or less often opposite; uppermost leaves scaly. Capitula terminal 
on stem and flowering shoots, two to six, narrowly-cylindrical, 1.8-2.8 
cm long (including ligulate florets), up to 3.5 cm long at fruiting, 3-7 
mm wide. 12-15-flowered, ellipsoidal in bud. Involucre glabrous or 
slightly pubescent, of 10-12 bracts; outermost bracts similar to upper 
cauline leaves with more or less wide scarious margin; outer bracts ovate; 
inner ones linear-oblong, somewhat acuminate. Ligulate florets 
yellow, pinkish on drying, less often white. Achenes 7 mm long, ribbed, 
angular, slightly hairy at apex, rest glabrous; pappus snow-white, three to 
four times as long as achene, its bristles plumose, barbed-scabrous only 
in upper part. Flowering July to August. 



81 

Saline meadows, solanchaks, saline sands, loess with salty coating. — 
Soviet Central Asia: Balkhash Region (Hi River Basin). General 
disribution: Dzhungaria-Kashgaria, Mongolia, China. Described from 
Mongolia. Type in Leningrad. 

Note. A highly variable species, representing the Mongolian element 
in the flora of the USSR. I have not seen the type of S. fengtiensis Nakai, 
a species described from northeastern China, but on the basis of the 
description, 1 have included it, with a question mark, among the synonyms 
of S. mongolica Maxim. 

Section 9. Turkestanicae Lipsch. in Addenda XXVIII, 720. — Stems 
erect, usually highly branched, leafy. Leaves linear or broadly lanceolate. 
Capitula usually in racemes, many-flowered or less often few-flowered. 
Involucre many-rowed. Ligulate florets yellow or orange. Achenes 
glabrous, cylindrical, ribbed, smooth. Pappus dirty white, wholly 
deciduous, its bristles plumose, 5-10 of them longer, barbed above. 

Series 19. Franchetianae Lipsch. in Addenda XXVIII, 721. — Basal 
and cauline leaves wide, ovate-elliptical or broadly lanceolate. 

46. S. franchetii Lipsch. in Byull. Mosk. Obshch. Isp. Pr. Otd. Biol. 
XLII, 2 (1933) 154; Lipsch. Fragm. Monogr Scorzonera II (1939) 150.— 
S. racemosa auct. non Franch.: Lipsch. Sist. Zam. ob Aziat. Skorts. (1932) 
4, p. min. p.; Pavl. in Tr. po Prikl. Bot. Ser. I, 2 (1937) 288, p. p.— 5. 
macrophylla B. Fedtsch. ex Pavl. op. cit. (1937) 289, non Fisch. and 
Mey.— Ic: Lipsch. op. cit. (1939) Plate 26. 

Perennial. Root cylindrical, vertical; root collar slightly covered with 
indurescent brown sheaths of basal leaves. Stems solitary or several, 40- 
70 cm high, erect, glabrous or sparsely pubescent, sulcate, leafy, branched 
above, branches 3-10(20) cm long, each terminating in capitulum. Basal 
and lower cauline leaves long-petiolate, hard, ovate-elliptical to broadly 
lanceolate, (13.5)20-30(35) cm long (including petioles) and (1.5)2-3(8) 
cm wide, acuminate, along margin flat or slightly undulate, serrulate under 
hand lens; venation almost pinnate, midrib usually wider, prominent, 
usually broadened at junction with petiole, carinate beneath, lateral veins 
not prominent; middle cauline leaves linear-lanceolate or lanceolate, 
(6.5)10-15 cm long and 0.5-2 cm wide; upper leaves reduced, 1.5-4.5 
cm long. Capitula (3)7-15 on single stem, broadly cylindrical, 2.5-3.5 
cm long (including ligulate florets), terminal on branches. Involucral bracts 
82 many-rowed, rigid, pubescent, glabrescent; outer bracts small, deltoid; 
middle bracts ovate; inner bracts lanceolate, sometimes with somewhat 
carinate midrib, obtuse, scarious along margin. Ligulate florets yellow, 
sometimes slightly pinkish when dry. Achenes 0.7-1.2 cm long. 



82 

cylindrical, glabrous, smooth, ribbed; pappus dirty-white, wholly 
deciduous, pappus bristles soft, plumose, five of them longer, plumose at 
base, barbed, scabrous above. Flowering July to August. 

Meadow-covered slopes, sometimes as weed in crop fields. — Soviet 
Central Asia: Syr-Darya (Karatau). Endemic. Described from Syr-Darya 
Karatau. Type in Moscow. 

Series 20. Pauciflorae Lipsch. in Addenda XXVIII, 72 1 .— Capitula 
narrowly cylindrical, few-flowered. Involucral bracts 12-15. 

47. S. turkestanica Franch. in Ann. Sc. Nat., ser. VI, Bot. XVI (1883) 
333; Lipsch. Sist. Zam. ob Aziat. Skorts. (1932) 3, 4-6; Lipsch. Fragm. 
Monogr. Scorzonera II (1939) 142. — S. nuristanica Kitam. in Acta 
Phytotax. et Geobot. XVII (1957) 39 and Fl. Afghan. (1960) 443 
Koalevskaja in Fl. Uzb. VI, 439 (excl. syn. S. racemosa Franch.). — Ic. 
Lipsch. op. cit. (1932) 25, fig. 1; 27, fig. 3; op. cit. (1939) Plate 22 
Kitam. op. cit. (1960) fig. 103. 

Perennial. Root vertical, slender; root collar sparsely covered with 
brown or yellow leaf sheaths. Stems 30-80 cm high, solitary or several, 
erect, somewhat sulcate, weakly pubescent or almost glabrous, leafy. 
Leaves sparsely pubescent or almost glabrous; basal and lower cauline 
leaves linear or linear-lanceolate, (2)3-9(12) mm wide, with three to five 
veins and distinct, whitish midrib sometimes keeled; leaves at base 
narrowed into long sulcate petiole; middle cauline leaves sessile, slightly 
amplexicaul, linear, flat, acuminate, gradually reduced upward. Capitula 
in racemes almost from middle of stem, narrowly cylindrical, 2-3.2 cm 
long and 0.4-0.6 cm wide, remote, solitary (less often in twos and threes) 
on short peduncles (longer in many specimens). Capitula 6-8(12)- 
flowered, ellipsoidal or cylindrical in bud; involucre puberulent, 
glabrescent, bracts 12-15, unequal; outer bracts deltoid, acuminate, with 
indistinct midrib; middle bracts ovate; inner ones linear-lanceolate, two 
to three times as long as outer. Ligulate florets yellow. Achenes 10- 
13(20) mm long, glabrous, smooth, ribbed; pappus dirty-white, its bristles 
plumose, 10 of them longer, barbed above. Flowering July to August. 

Turfy slopes, meadows, herb steppe, forest glades, grassy parts 
83 of juniper forests. — Soviet Central Asia: Syr-Darya, Tien Shan, Pamiro- 
Alai (northern part). Endemic. Described from western Tien Shan. Type 
in Paris; isotype in Leningrad. 

Note. I did not see the type of S. nuristanica Kitam. (a species 
described from Nuristan; the type is preserved in the herbarium of Kyoto 
University). However, the figure of S. nuristanica Kitam. leaves no doubt 
about its identity with S. turkestanica Franch. Therefore, I include it among 



83 

the synonyms of the latter. S. turkestanica Franch. evidently remained 
unknown to Kitamura, the author of the species, as he mistakenly compares 
his 5. nuristanica with S. purpurea. 

Series 21. Racemosae Lipsch. in Addenda XXVIII, 721. — Capitula 
broadly cylindrical, many-flowered. Involucral bracts 20-32. 

48. S. iliensis Krasch. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 1 
(1933) 178; Kovalevskaja in Fl. Uzb. VI, 439. — S. racemosa auct. non 
Fisch.: Lipsch. Sist. Zam. ob Aziat. Skorts. (1932) 3, 4, p. max. p.; Pavl. 
in Tr. Prikl. Bot. Ser. I, 2 (1937) 288, p. p.; Lipsch. Fragm. Monogr. 
Scorzonera II (1939) 143.— Ic: Lipsch. op. cit. (1939) Plate 23. 

Perennial. Root vertical. Stem (25)35-70(90) cm high, solitary or 
several, erect, distinctly sulcate, glabrous, or less often somewhat floccose, 
glabrescent, more or less branched or very rarely simple. Basal leaves 
linear, (2)4-10 mm wide, narrowed into long, sulcate petioles, base 
broadended into membranous sheaths loosely covering root collar; leaves 
hard, flat or folded, scabrous along margin, with three-five(seven) veins; 
venation parallel, with somewhat prominent veins, midrib usually white, 
often carinate, broadened toward petiole; cauline leaves narrow-linear, 
sessile, slightly amplexicaul, arcuately divergent, reduced above. Capitula 
on peduncles, more or less numerous, less often solitary, forming indistinct 
corymbose head; peduncles with reduced leaves; capitula globular in bud. 
Involucre arachnoid-pubescent, many-rowed, glabrescent, bracts rigid, 20- 
32, slightly membranous along margin; outermost bracts small, deltoid, 
those in second row ovate-deltoid, inner ones lanceolate, obtuse, with or 
without prominent carinate midrib. Ligulate florets yellow, strongly 
exserted from involucre. Achenes up to 15 mm long, glabrous, smooth, 
ribbed; pappus dirty white, its bristles plumose, 5-10 of them longer, 
barbed above. Flowering June to August. 

Dry stony slopes, meadow slopes, among shrubs, saline meadows on 
floodplains. — Soviet Central Asia: Balkhash Region (Hi River valley), Syr- 
Darya (Karatau), Tien Shan (western), Dzhungaria-Tarbagatai. Endemic. 
Described from Hi River. Type in Leningrad. 

Note. Acquaintance with the photograph of the type of S. racemosa 
Franch. — the species described from Namangan in Fergana (type preserved 
in Paris) — has shown me that earlier I had wrongly identified it with 5. 
84 iliensis Krasch. The present condition of the type of S. racemosa Franch. 
did not permit me to clarify the taxonomic position of S. racemosa Franch. 
(non Lipsch.) in the classification of the genus. This species remains 
unclear to me. Kovalevskaja {Flora of Uzbekistan, VI (1962) 439) included 
5. racemosa Franch. among the synonyms of S. turkestanica Franch. 



84 

49. S. transiliensis M. Pop. in Lipsch. Fragm. Monogr. Scorzonera 
II (1939) 148. — S. purpurea var. subincana Rgl. and Herd, in Bull. Soc. 
Nat. Mosc. XL, 3 (1867) 170. — S. racemosa auct. non Franch.: Lipsch. 
Sistem. Zam. ob Aziat. Skorts. (1932) 4, p. min. p. — Ic: Lipsch. op. cit. 
(1939) Plates 24 and 25. — Exs.: Lipsch. Herb, cum sched. lithogr. a 
CUNIPMED edit. (1933) No. 200 (sub nom. S. racemosa Franch.). 

Perennial. Root vertical, producing one to three shoots and sometimes 
rosettes of basal leaves; root collar sparsely covered with leaf sheaths. 
Stems 25-75 cm high, indistinctly sulcate, glabrous or weakly floccose, 
simple or often branched above, branches two or three (five), 3.5-12 cm 
long, each with single capitulum. Basal and lower cauHne leaves simple, 
Unear or less often linear-lanceolate, 0.4-0.8(1) cm wide, acuminate, along 
margin scabrous and serrulate (hand lens!), puberulent, veins narrow, 
parallel lamina (especially in basal leaves) narrowed at base into long 
petiole; upper cauline leaves and those on branches reduced, narrowly 
linear, subulate. Capitula cylindrical, 3-3.5 cm long, one to three(five). 
Involucre of rigid bracts, in three to five rows; outer bracts smaller, deltoid- 
ovate; middle bracts lanceolate; inner ones lanceolate or linear-lanceolate, 
obtuse; all bracts arachnoid-pubescent, glabrescent. Ligulate florets dark 
orange, about 1.5 cm long and 2.5 mm wide, five-toothed at tip. Achenes 
narrowly terete, 1-1.5 cm long, glabrous, slightly ribbed, whitish, reddish 
when mature; pappus dirty white, wholly deciduous, its bristles plumose, 
five of them longer, barbed. Flowering July to August. 

Meadows on mountain slopes, secondary herb meadows in the 
Schrenk spruce {Picea schrenkiana Fisch. and Mey.) zone, often on clayey 
slopes. — Soviet Central Asia: Tien Shan (Trans-Ili Alatau, Ketmen 
Mountains, Kungei Alatau). Endemic. Described from Trans-Ili Alatau. 
Type in Alma Ata. 

Note. S. transiliensis M. Pop. is a race quite close to S. iliensis 
Krasch., from which it differs at once when flowering by its dark orange 
Ugulate florets, which are very conspicuous. Variation in leaf width is 
observed, particularly, in herbarium collections from the region of the 
Sardzhass Valley (eastern Tien Shan), which are distinguished by narrow 
leaves (1-2 mm wide) and greater xerophily and are being recognized as 
a separate variety — var. angustifolia Lipsch. (op. cit. (1939) 150). This 
species was erroneously identified by earlier workers (Regel, Herder) as 
5. purpurea, with which it has nothing in common. 

Section 10. Foliosae (Boiss.) Lipsch. comb, nova in Addenda XXVIII, 
721.— Subsection Foliosae Boiss. Fl. or. Ill (1875) 756.— Plant pubescent 
85 or glabrous. Stems leafy, often tall. Outer achenes more or less tuberculate 
or scaly-acerate along ribs. 

Series 22. Strictae Lipsch. in Addenda XXVIII, 721. 



85 

50. S. gracilis Lipsch. in Addenda XXVIII, 721. 

Perennial. Whole plant grayish, densely fine-tomentose from stellate 
hairs. Root cylindrical; root collar covered with broadened brown 
indurescent sheaths of previous year's basal leaves and current year's 
hyaline, weakly glossy leaf sheaths. Stems 15-20 cm high, erect or slightly 
ascending, simple or sparsely branched. Leaves narrowly linear, densely 
pubescent; basal leaves 2-4 mm wide, often folded lengthwise, subulate 
at apex, almost as long as stem or one-third shorter, base broadened into 
hyaline, glabrous sheath; cauline leaves reduced, few, sessile, slightly 
semiamplexicaul; solitary uppermost leaf reduced, often approximate to 
capitulum, forming bract with attenuate-aristate apex. Capitula solitary or 
two to five, large, 2.5-3 cm long, 7-8 mm wide (at flowering). Involucre 
canescent; outer bracts ovate or deltoid-ovate; inner ones oblong- 
lanceolate; all bracts more or less acuminate, membranous along margin, 
especially inner ones. Ligulate florets pink, considerably longer than 
involucre (two times as long). Mature achenes unknown, young achenes 
ribbed, acerate along ribs; pappus of unequal bristles, bristles plumose, 
barbed-scabrous above. Flowering May. 

Stony slopes. — Soviet Central Asia: Pamiro-Alai (Shugnan). Endemic. 
Described from vicinity of Khorog. Type in Leningrad. 

Note. It is easily distinguished from the sister species S. pubescens 
DC. by its white-tomentose pubescence and pink flowers. 

51. S. stricta Homem. Hort. Hafn. II (1813) 750; M.B. Fl. taur.- 
cauc. Ill (1819) 522; DC. Prodr. VII, 1 (1838) 122, p. p.; Boiss. Fl. or. 
Ill, 768, excl. syn.; Schmalh. H. II, 142; Krasch. in Fl. Yugo-Vost. VI, 
448; Grossh. Opred. Rast. Kavk. 511; Kryl. Fl. Zap. Sib. XI, 2991.— 
5. villosa M.B. op. cit. II (1808) 235, excl. syn. — 5. marschalliana auct. 
non C.A.M.: Ldb. Fl. Ross. II, 2 (1845-1846) 795, ex parte.— 5. hispanica 
angustifolia auct. Fl. Ross. — S. marschalliana auct. plur. fl. Ross, non 
C.A.M. 

Perennial. Plants more or less pubescent, greenish. Root cylindrical, 
up to 2.5 cm thick; root collar covered with light brown, entire or fibrous 
remnants of sheaths. Stems solitary, erect, 20-50 cm high, branched, 
branches 8-20 cm long, erect, flexuous. Leaves long, linear or linear- 
lanceolate, gradually narrowed toward tip, acuminate-subulate, flat or 
86 folded, 2-7 mm wide, with prominent keel beneath; basal leaves with 
broadened base; cauline leaves semiamplexicaul. Stem leafy throughout, 
particularly more densely in lower half. Capitula (2)3-10(20), on upright 
peduncles, cylindrical, 2.5-3(3.5) cm long and 5-7 mm wide (at 
flowering). Involucre arachnoid-pubescent; outer involucral bracts broadly 
lanceolate, usually with distinct, somewhat carinate midrib, acuminate; 
inner bracts oblong, acuminate. Ligulate florets yellow, longer than 



86 

involucre. Achenes 10-12 mm long, glabrous, with ring of hairs at tip at 
articulation of pappus; outer achenes tuberculate along ribs, more or less 
acerate; pappus dirty white, its bristles plumose, longer of them barbed- 
scabrous above. Flowering June to July. 

Steppes (meadow; sandy; feathergrass-sheep fescue-wormwood; with 
stands of Spiraea; Festuca; saline and Stipa lessingiana) and meadows. — 
European Part: Crimea, Bessarabia, Black Sea Region, Lower Don, Lower 
Volga, Trans- Volga, Volga-Don, Volga Kama; Western Siberia:. Upper 
Tobol, Irtysh (western part); Soviet Central Asia: Aralo-Caspian (northern 
part); Caucasus: Ciscaucasia, Western Transcaucasia, Dagestan. Described 
from ? Type in Copenhagen. 

52. S. pubescens DC. Prodr. VII, 1 (1838) 122; Kryl. Fl. Zap. Sib. 
XI, 2990.— 5. stricta auct. non Homem.: Ldb. Fl. alt. IV (1833) 163; 
Kryl. Fl. Alt. Ill (1904) 735.-5. stricta auct. fl. turkest. p. max. p.— 5. 
marschalliana C.A.M. in Mem. Ac. Petersb. VI, ser. Sc. nat. IV (1841) 
200. 

Perennial. Plant grayish from short-branched hairs. Root cylindrical, 
vertical; root collar covered with brown membranous leaf sheaths. Stems 
solitary or several, (3.5)7-20(35) cm high, more or less branched, often 
from base, densely leafy below. Leaves 5-20 cm long, linear, 1-5 mm 
wide, more or less curved, flat or folded, carinate beneath, margin 
sometimes undulate; basal leaves broadened into glossy sheath; cauline 
leaves somewhat broadened at base, semiamplexicaul, uppermost leaves 
reduced, subulate. Capitula solitary, terminal on stem and branches, 2- 
2.5 cm long and 3-7 mm wide (at flowering). Involucre pubescent, 
glabrescent; outer involucral bracts ovate-deltoid, usually with prominent 
midrib, sometimes slightly carinate; inner bracts broadly lanceolate, 
scarious along margin, both acuminate. Ligulate florets yellow, sometimes 
reddish below; outer florets one and one-half times as long as involucre. 
Achenes terete, (8)10-12(14) mm long, glabrous, outer achenes ribbed 
above, acutely or obtusely tuberculate: tubercles variable; pappus dirty 
white, longer than achenes, with crown of hairs at articulation of pappus, 
plumose, longer bristles barbed-scabrous above. Flowering May to June. 

Stony and gravelly steppe slopes of hills and mountains, rocks. — 
Western Siberia: Irtysh, Altai; Soviet Cenral Asia: Aralo-Caspian, Balkhash 
Region, Syr-Darya, Tien Shan, Dzhungaria-Tarbagatai. Endenuc. Described 
from Siberia. Type in Geneva. 
87 Note. Where the ranges of 5. pubescens and S. stricta meet in western 

Kazakhstan, intermediates between the two species are found. 

Series 23. Hispanicae Lipsch. in Addenda XXVIII, 721. — Leaves 
ovate-lanceolate or lanceolate. 



87 

53. S. hispanica L. Sp. pi. (1753) 791; Willd. Sp. pi. Ill, 3 (1803) 
1499, p. p.; DC. Prodr. VII, 1 (1838) 120, p. p.; Ldb. Fl. Ross. II, 2, 794; 
Boiss. Fl. or. Ill, 767; Grossh. Fl. Kavk. IV (1934) 241; Grossh. Opred. 
Rast. Kavk. 511.— 5. taurica M.B. H. taur.-cauc. II (1808) 234; III (1819) 
521; DC. Prodr. VII, 1 (1838) 121; Stank, and Tal. Opred. Vyssh. Rast. 
(1949) 687; Isaev in Fl. Azerb. VIII, 523.-5. glastifolia Willd. op. cit. 
(1803) 1499.-5. hispanica^. glastifolia (Willd.) Wallroth. Ann. hot. 
suppl. Fl. gall. (1815) 95. — 5. crispatula auct. non Boiss.: Stank, in Stank, 
and Tal. op. cit. (1949) 687. 

Perennial. Plant pubescent, glabrescent. Root cylindrical. Stems 
(25)35-50(75) cm high, erect, usually branched in upper part or from 
middle; branches erect; stem base covered with more or less numerous, 
broadened, indurescent, glossy, stramineous or brown sheaths of basal 
leaves. Leaves crowded in lower third of stem, ovate-lanceolate or oblong, 
with many veins (midrib often broader, hyaline), pubescent or glabrous, 
margin flat or slightly undulate, scabrous-toothed; lower leaves long- 
petiolate; middle leaves oblong-lanceolate, amplexicaul, acuminate; 
uppermost leaves subulate. Capitula few, terminal on branches, cylindrical, 
large, 2-2.5(3) cm long (at flowering). Involucre pubescent, glabrescent; 
outer involucral bracts ovate; inner oblong-lanceolate, all bracts acuminate. 
Ligulate florets yellow, one and one-half times as long as involucre. 
Achenes (mainly outer) 10-14 mm long, angular-ribbed, more or less 
finely rugose-tuberculate along ribs; pappus dirty- white, its bristles 
plumose, longer of them barbed-scabrous above. Flowering May to June. 

Meadows, steppes. — European Part: Volga-Middle Dnieper, Volga- 
Don, Trans- Volga Region, Lower Volga, Black Sea Region, Crimea; 
Western Siberia: Upper Tobol; Caucasus: Ciscaucasia, Eastern and 
Western Transcaucasia, Dagestan; Soviet Central Asia: Aralo-Caspian 
Region (northern part). Type in London. 

Note. 5. hispanica L. was described from cultivated plants. Probably, 
the name 5. taurica M.B. is more appropriate for our representatives of 
this species aggregate. However, following tradition, pending a 
monographic study of 5. hispanica s. 1., I am tentatively placing this 
plant under the widely used name 5. hispanica L. 

54. S. inconspicua Lipsch. ex Pavl. in ByuU. Mosk. Obsch. Isp. Prir. 
Otd. Biol., XLII, 2 (1933) 139; Popov in Tr. Uzb. Gos. Univ. Nov. Ser. 
27, Biol. 14 (1941) 92; Kryl. Fl. Zap. Sib. XI, 2992; Kovalevskaja in Fl. 
Uzb. VI, 440. — 5. marschalliana var. oblongifolia Trautv. in Bull. Soc. 
Nat. Mosc. 39, 2 (1866) 383. — 5. marschalliana var. latifolia Rupr. in 
Mem. Acad. Sc. Petersb. VII, ser. XIV, 4 (1869) 58.-5. taurica auct. fl. 
turk., non M.B. — 5. hispanica auct. fl. turkest. pro max. parte, non L. 
nee Willd. — 5. macrophylla Herd, in herb. — Ic: Pavlov in Byull. Mosk. 



Obshch. Isp. Prir. Otd. Biol. XLII, 2 (1933) 139.— Exs.: Pavl. and Lipsch. 
PI. exs. edit. CUNIPMED (1932) No. 50. 

Perennial. Plant weakly pubescent, sometimes glabrescent. Root collar 
covered with indurescent brown or stramineous sheaths of basal leaves. 
Stems 10-25(35) cm high, pubescent, mostly branched from base or 
middle; branches curved, flexuous, terminating in solitary capitula. Basal 
leaves broadly lanceolate, hard, curved, with many veins, margin 
cartilaginous, usually undulate and scabrous, pubescent with branched hairs 
or almost glabrous, narrowed at base into petiole; cauline leaves sessile, 
semiamplexicaul, narrower than basal leaves, undulate along margin, 
gradually reduced above. Capitula 2-^(10), long cylindrical, 2-2.5(3) cm 
long and 0.5-0.8(1) cm wide (at flowering), broader at fruiting, terminal 
on branches. Involucre pubescent; bracts coriaceous, outer bracts deltoid- 
ovate, somewhat acuminate; inner bracts oblong-lanceolate, narrowly 
membranous along margin, more or less acuminate or subobtuse, several 
times as long as outer. Ligulate florets yellow, one and one-half to two 
times as long as involucre. Mature achenes terete, 10-15 mm long, 
glabrous, ribbed, somewhat acerate along ribs, sometimes only in upper 
part scabrous; pappus dirty white, its bristles plumose, longer of them 
barbed above; crown of hairs at articulation of pappus with achene. 
Flowering May to July. 

Stony-gravelly slopes. — Western Siberia: Irtysh; Soviet Central Asia: 
Syr-Darya, Tien Shan, Pamiro-Alai, Balkhash Region, Dzhungaria- 
Tarbagatai. Endemic. Described from Syr-Darya Karatau. Type in 
Moscow; isotype in Leningrad. 

Note. The Pamir specimens differ in having dense grayish-pubescent, 
scapelike, and shorter (7-10 cm high) stems. Evidently, it is a separate 
race, needing observations in the field and new, more complete collections 
(var. incana Lipsch.). — Collections from Kirgizia (vicinity of Lake Issyk- 
Kul, Ortotokoi, on mesas V. 1941. O.E. Knorring) differ from the type 
as follows: plant green, almost glabrous; capitulum smaller; involucral 
bracts narrower, all lanceolate; inner bracts with distinct veins; mature 
outer achenes 13 mm long, olive-colored, covered with spinules along 
entire length. Apparently, it is a separate species, but the available material 
is insufficient to describe it. — 5. inconspicua Lipsch. occurs mixed with 
S. pubescens DC. in western Kazakhstan. 

55. S. chantavica Pavl. in Vestn. Akad. Nauk Kazakh. SSR 8 (1950) 
27.— Ic: Pavl. op. cit. 28, Fig. 19. 
89 Perennial. Plant slightly pubescent. Root woody, thickened at apex, 

many-headed; root collar covered with indurescent remnants of petioles 
of basal leaves. Stems 25-30 cm high, slender, sulcate, simple or branched. 
Basal leaves rosulate, obovate-lanceolate, 8-10 cm long, 1.5-2 cm wide. 



89 

narrowed into broad-winged petiole, grooved on upper side; leaves 
broadest in middle, somewhat acuminate, cartilaginous along margin, 
slighdy undulate; cauline leaves reduced, lanceolate, sessile, somewhat 
amplexicaul. Capitula on long peduncles, cylindrical, 2.5 cm long, 0.6- 
0.7 cm wide (at flowering). Involucre arachnoid-pubescent, glabrescent; 
outer bracts broadly ovate or roundish, 5-6.5 mm long, short-acuminate; 
middle bracts oblong, broadly lanceolate, acuminate, 10-11 mm long; 
inner bracts oblong-lanceolate, 15-25 mm long, subobtuse, with red stripe 
along margin and narrowly membranous, with one vein. Ligulate florets 
yellow, exserted from involucre. Achenes cylindrical, mature achenes 
(10)12-13 mm long, brown, ribbed, short-scaly-acerate along ribs; pappus 
dirty white, as long as achene, its bristles plumose, longer of them barbed- 
scabrous above. Flowering May to June. 

Stony rubbly slopes. — Soviet Central Asia: Tien Shan (Chu-Ili 
Mountains). Endemic. Described from Chu-Ili Mountains. Type in Alma- 
Ata. 

Section 11. Polycladae DC. Prodr. VII, 1 (1838) 125; Benth. and 
Hook. f. Gen. pi. pi. II (1873) 532; O. Hoffm. in Pflanzenfam. IV, 5 
(1893) 365; Lipsch. Fragm. Monogr. Scorzonera II (1939) 1 1 1 .—Capitula 
few-flowered, comparatively small. Stems highly branched, branches 
mostly virgate, sparsely leafy. 

Series 24. Acanthocladae Lipsch. in Addenda XXVIII, 722. — 
Semishrubs, spherical in habit, usually with numerous prickly remnants 
of previous year's branches. Stems with number of partly prickly sterile 
branches. Cauline leaves small, linear-subulate. Capitula narrowly 
cylindrical, few-flowered. 

56. S. acanthoclada Franch. in Ann. Sc. Nat. ser. VI, XVI (1883) 
333; Lipsch. Sist. Zam. ob Aziat. Skorts. (1932) 8; Tekutiev in Zhum. 
"Sov. Kauchuk" 3 (1933) 37; Vel'tishchev and Vel'tishcheva in Zhum. 
"Pripoda", 9 (1934) 62; Lipsch. Fragm. Monogr. Scorzonera II (1939) 
113; Popov in Tr. Uzb. Cos. Univ. Nov. Ser. 27, Biol. 14 (1941) 94; 
Lipsch. in Fl. Turkm. VII, 308; Kovalevskaja in Fl. Uzb. VI, 440.— 5. 
chondrilloides Rgl. and Schmalh. in Izv. Obshch. Lyubit. Estestv., Antrop. 
i E'tnogr. XXXIV, 2 (1882) 54 (non S. chondrilloides Pourr. ex Willd., 
1803).— Ic: Zhurn. "Priroda", 9 (1934) pp. 62-64; Lipsch. op. cit. (1939) 
plate 17. 

Perennial. Plant weakly pubescent, glaucescent semishrub, often 
spherical in habit. Root cordlike, with several small lateral roots, growing 
deep into soil, often twisted, producing numerous caudices near root collar, 
forming turf, covered with dark brown remnants of previous year's leaf 



90 

90 sheaths. Stems usually numerous, slightly curved, dichotomously branched, 
successively branched forming slightly spinose branches of second and 
third order; previous year's (dry) stems usually with numerous lateral 
spinescent branches. Basal leaves narrowly linear or linear-lanceolate, 
mostly with three veins, almost as long as stem; cauline leaves reduced, 
linear-subulate. Capitula narrowly cylindrical, about 1.5-2.5 cm long and 
2-8 mm wide, on somewhat long flowering branches (isolated specimens 
having up to 1,500 capitula!), with three to five flowers. Involucre of 
few appressed pubescent bracts. Ligulate florets yellow, red on drying. 
Achenes 10-12 mm long, ribbed, smooth, few in each capitulum; pappus 
dirty white, as long as achene, pappus bristles plumose to two-thirds, 
barbed-scabrous above, with ring of hairs at articulation of pappus with 
achene. Flowering July to August. 

Slopes of northern, northwestern, and northeastern exposure within 
altitudinal Umits of 2,200-3,800 m, in alpine and subalpine zones. On 
clayey slopes, forming part of steppe-type cenoses with a predominance 
of grasses; on slopes with coarse poor soils, forming groupings with 
predominance of montane xerophytes. — Soviet Central Asia: Pamiro-Alai. 
General distribution: Afghanistan. S. acanthoclada Franch. described from 
Archamaidan. Type in Paris. 

Note. S. chondrilloides Rgl. and Schmalh. was described from Central 
Asia (Kekh); the type is preserved in Leningrad. 

57. S. kuhistanica M. Pop. in Tr. Uzb. Cos. Univ. Nov. Ser. 27, 
Biol. 14 (1941) 94. 

Similar to preceding species. Branches not spinescent. Stems 20-30 
cm high. All leaves linear; basal leaves up to 30 cm long, capitula not 
large, at flowering 1.2-1.5 cm and at fruiting 2.5 cm long. Achenes 14 
mm long, with violet stripes. 

On slopes in the glacier region. — Soviet Central Asia: Pamiro-Alai 
(Kugistan). Endemic. Described from Zeravshan (Kugistan). Type in 
Tashkent. 

Note. I did not see the original specimens of this species, which the 
author himself says is connected by intermediates with 5. acanthoclada 
Franch. I think that 5. kuhistanica M. Pop. should include 5. acanthoclada 
var, gracilis Tekut., which was characterized by Tekutiev by the following 
characters: "Plant taller (25-70) cm high); stems fewer (five to eight), 
less branched, comparatively little curved; leaves glabrous even when 
young, linear-lanceolate, wider than in the typical form (3-8 mm wide), 
basal leaves with four to six veins; capitula five- to eight-flowered (less 
often with 12 flowers)." Distributed in Darvaz and Shugnan. In order to 
clarify the relationship between the species pair 5. kuhistanica — S. 
acanthoclada, new observations and collections from the field are needed. 



91 

I did not find a single specimen of S. acanthoclada without prickly 
branches in the Herbarium of the Botanical Institute, Academy of Sciences 
of the USSR, which could be identified as S. kuhistanica. Kovalevskaja 
91 {Fl Uzb. VI (1962) 440) included 5. kuhistanica M. Pop. among the 
synonyms of S. acanthoclada Franch. Tentatively, I have left the former 
in the key as a separate taxon, in order to attract the attention of 
investigators to this confusing species. 

In Symb. Afgh. II (1955) 195 and Fig. 135, 137, a new species, S. 
polyclada Rech. f. and Koie, is described from Afghanistan, compared 
by the authors with S. tortuosissima Bouss., and referred to subsection 
(in my interpretation, section) Intricatae Boiss. I did not see either the 
types of this species (they are preserved in Vienna and Copenhagen) or 
the other original specimens. Judging from the description and figures, it 
is quite probable that S. polyclada Rech. f. and Koie represents 5. 
acanthoclada s. 1., more precisely, var. gracilis Tekut. (= S. kuhistanica 
M. Pop.). This is all the more probable because S. acanthoclada is not 
cited by Rechinger in his work on Afghanistan. 

Series 25. Divaricatae Lipsch. in Addenda XXVIII, 722. — Semishrubs 
or perennials without numerous prickly remnants of previous year's stems. 
Stems without sterile branches. Cauline leaves well developed. 

58. S. pseudodivaricata Lipsch. in ByuU. Mosk. Obshch. Isp. Prir. 
Otd. Biol. XLII, 2 (1933) 158; Kotov in Sov. Bot. 3 (1935) 68, 69; Lipsch. 
Fragm. Monogr. Scorzonera II (1939) 128. — S. divaricata var. foliata 
Maxim, in Bull. Acad. Sc. Petersb. XXXII, 4 (1888) 494.-5. divaricata 
var. virgata Maxim, op. cit. (1888) 495. — Ic: Lipsch. op. cit. (1939) 
Plates 19 and 20. 

Perennial. Semishrub with few caudices. Root collar covered with 
indurescent, entire, or fibrous sheaths of basal leaves. Stems 10-45 cm 
high, numerous, glabrous, or pubescent, subsequently woody, virgate, 
branched always from middle or above. Basal leaves up to 17 cm long, 
broadened at base into brown or stramineous sheaths; cauline leaves 
alternate, sometimes almost opposite at base of branches, somewhat falcate, 
narrowly linear, 1-9 cm long and (0.5)2-4.5 mm wide, acuminate, 
sometimes uncinate; upper cauline leaves reduced to scales. Inflorescence 
lax corymbose panicle. Capitula usually numerous, narrowly cylindrical, 
1.3-2.5 cm long (including ligulate florets), solitary, terminal on branches, 
7-12-flowered. Involucral bracts many-rowed, glabrous or pubescent; outer 
bracts small, triangular, somewhat acuminate; middle bracts ovate, two to 
three times as long; inner bracts oblong-lanceolate, almost two times as 
long as middle ones. Achenes 5-10 mm long, terete, sometimes slightly 
curved, yellow or dark green, glabrous or with ring of hairs below pappus, 



92 

ribbed, ribs turberculate or smooth; pappus white, its bristles plumose, 
barbed above. Flowering July to August. 
92 Rocks, stony screes, outcrops of marblelike limestone and quartzites, 

pebbles. — Soviet Central Asia: Tien Shan (central Angiz area). General 
distribution: Dzhungaria-Kashgaria, Mongolia, China. Described from 
Mongolia (Hangai). Type in Leningrad. 

Section 12. Pusillae Lipsch. in Lipsch. Fragm. Monogr. Scorzonera 
II (1939) 59. — More or less spreading perennials with tuberous roots. 
Leaves twisted at apex. Capitulum narrowly cylindrical, usually numerous 
(up to 50). Ligulate florets slightly exserted from involucre, yellow, red 
on drying. Achenes narrow, terete or prismatic, with longitudinal ribs, 
glabrous, smooth, or less often slightly scabrous. 

59. S. pusilla Pall. Reise II (1773) 329 and Anh. 744; Willd. Sp. pi. 
Ill, 3, 1501; M.B. Fl. taur.-cauc. II, 237; Eichwald, PI. nov. (1831-1833) 
2, descr. emend.; Ldb. Fl. Alt. IV, 160; DC. Prodr. VII, 1, 118; Ldb. Fl. 
Ross. II, 2, 791; Boiss. Fl. or. Ill, 771; Maxim, in Bull. Acad. Sc. Petersb. 
XXXII, 4 (1888) 492; Paczoski, Florograf. i Fitogeogr. Issl. Kalmytsk. 
Step. (1892) 86; Schmalh. Fl. II, 144; Grossh. Fl. Kavk. IV (1934) 239; 
Krasch. in Fl. Yugo-Vost. VI, 451; Lipsch. Fragm. Monogr. Scorzonera 
II (1939) 59; Kryl. Fl. Zap. Sib. XI, 2986; Isaev in Fl. Azerb. VIII, 524; 
Lipsch. in Fl. Turkm. VII, 308; Kovalevskaja in Fl. Uzb. VI, 443.-5. 
circinnata Pall. Reise III (1776) 677. — S. astrachanica DC. Prodr. VII, 1 
(1838) 118; Paczoski, op. cit. (1892) 87.-5. scoparia Claus ex Lipsch. 
op. cit. (1939) 61, non Boiss. and Kotschy, nomen. — Dschangiiria 
desertorum Less, ex Lipsch. op. cit. (1939) 61, nomen. — S. popovii Lipsch. 
Fragm. Monogr. Scorzonera I (1935) 23.— Ic: Pall. Reise II, 2 (1773) 
tab. L; Lipsch. op. cit. (1939) Plate 8. 

Perennial. Plant (4)10-25(40) cm high, mostly spreading. Roots with 
single, sometimes many tubers; root collar covered with sheaths, sheaths 
subsequently fibrous, often pubescent on inner side. Stems flexuous, 
sulcate, mostly branched, less often simple, pubescent (sometimes floccose) 
or glabrous, leafy. Leaves alternate, less often opposite or even almost 
whorled (in groups of three), mostly arcuately bent, glaucous, pubescent, 
narrowly linear, 1-1.5(2) mm to 3-4.5 mm wide (var. latifolia Lipsch.), 
with three veins, flat or folded, with filiform, uncinate or slightly twisted 
tip. Capitula narrowly cylindrical, 1-3 cm (including ligulate florets), at 
fruiting 1.7-4.5 cm long, on somewhat spreading peduncles, 5-20 (up to 
50!) per plant. Involucre of few bracts, arachnoid-pubescent; outer bracts 
reduced, ovate; middle ones oblong, obtuse; inner ones oblong-lanceolate 
or linear-lanceolate, scarious along margin, subobtuse. Ligulate florets 
yellow, pink when dry, one-fourth to one-third longer than involucre. 



93 

93 Achenes 7-12 mm long, glabrous, with longitudinal ribs; ribs smooth, 
sometimes some of them spinulose, resulting in scabrous achene; pappus 
white, two to two and one-half times as long as achene, pappus bristles 
spreading, plumose up to two-thirds length, barbed-scabrous above. 
Flowering (March) April to May (June). 

Saline lands, stony slopes, sands, sandy steppes. — European Part: 
Lower Volga; Western Siberia: Upper Tobol (southern part), Irtysh; 
Caucasus: Southern Transcaucasia (Nakhichevan); Soviet Central Asia: 
Kara-Kum, Kyzyl-Kum, Amu-Darya, Aralo-Caspian, Syr-Darya, Pamiro- 
Alai, Tien Shan, Dzhungaria-Tarbagatai. General distribution: Iran, 
Dhungaria-Kashgaria, Mongolia. Described from the Caspian desert. Type 
in London. 

Note. As reported by several travelers and collectors, the tubers of 
this species are edible. 

In a series of specimens, especially from Mangyshlak, Kyzyl-Kum, 
and eastern Kazakhstan the achenes are swollen, pyriform in the lower 
part. This is explained by the presence of a parasite inside the achene. 

Section 13. Papposae Lipsch. and Krasch. in Lipsch. Fragm. Monogr. 
Scorzonera I (1935) 45. — Plants mostly with thick, tuberous roots. Stems 
branched from middle or base. Basal leaves petiolate, broadly ovate or 
oblong-lanceolate. Stems prostrate, scapelike, leafy only at base and lower 
third, or tall and densely leafy. Capitula narrowly or broadly cylindrical. 
Involucral bracts coriaceous; outer bracts ovate-deltoid, obtuse; inner ones 
oblong-lanceolate. Achenes flattened-cylindrical, mostly white, ribbed, 
slightly curved, with transverse rows of scabrous tubercles. Pappus snow- 
white, its bristles soft, densely plumose, inner ones longer, thicker above, 
somewhat barbed. Ligulate florets yellow, pink, or (as a result of drying?) 
lilac colored. 

Series 26. Darvasicae Lipsch. in Addenda XXVIII, 722. — Stem tall, 
leafy to tip, intemodes long. Capitula narrowly cylindrical. 

60. S. tadshikorum Krasch. and Lipsch. in Lipsch. Fragm. Monogr. 
Scorzonera I (1935) 49.— Ic: Lipsch. op. cit. (1935) Plate 18. 

Perennial. Underground part and root collar not known; possibly, 
root reduced, a tuber. Stems 20-30 cm high, sulcate, slightly pubescent, 
dichotomously branched; branches (floral branches) simple or branched, 
8-12 cm long. Leaves lanceolate, (5.5)7-11(14) cm long, 0.7-1.5(2) cm 
wide, narrowed below, amplexicaul, at base roundish or somewhat keeled 
with flat or barely undulate-crimped margins, veins except white midrib, 
indistinct; upper leaves reduced, with axillary flowering shoots; intemodes 
long. Capitula narrowly cylindrical, 2-2.5 cm long. Involucre glabrous; 
bracts slightly membranous along margins; outer bracts small, deltoid- 



94 

ovate; inner bracts oblong-lanceolate, subobtuse, longer than outer ones. 
Achenes terete, 7 mm long, slightly curved, somewhat tuberculate-toothed 

94 along ribs; pappus snow-white, its bristles soft numerous, plumose, five 
inner bristles longer, barbed above. Flowering June. 

Mountain slopes. — Soviet Central Asia: Pamiro-Alai (Darvaz, 
Childara). Described from Childara. Type in Leningrad. 

Note. The species was described from insufficient material (a single 
herbarium sheet) and needs further study and observations in the field. 

Series 27. Fer ganicae Lipsch. in Addenda XXVIII, 722. — Stems 
scapelike, branched from middle; flowering branches numerous; leaves 
crowded at base of fork. Intemodes shorter. Capitula narrowly cylindrical. 

61. S. ferganica Krasch. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. I, 
1 (1933) 180; Lipsch. Fragm. Monogr. Scorzonera I (1935) 52.— Ic: 
Lipsch. op. cit. (1935) Plate 19. 

Perennial. Plant weakly pubescent, glabrescent. Root thickened into 
tuber; root collar covered with long stramineous or brown sheaths. Stems 
scapelike, (8)10-16 cm high, dichotomously branched from middle; lateral 
branches (floral) (3)6-25, simple or branched, (3)8-12(15) cm long, 
unequal, bent, spreading. Leaves quite variable in form and size, with 
several veins and prominent midrib; basal leaves broadly linear, with 
yellow shining sheaths surrounding lower part of stem below branches 
and evidently embedded into stonty soil; cauline leaves linear-lanceolate 
or lanceolate, more or less acuminate, orbicular or cuneate at base, 
amplexicaul; margin undulate-crimped, (3)4—8(10) cm long and 0.4-1.5(2) 
cm wide. Achenes narrowly terete, (2)2.5-3 cm long and 0.4-0.6 cm 
wide. Involucre glabrous, involucral bracts with narrow membranous 
margin; outer bracts small, ovate-deltoid, 4-6 mm long, obtuse; inner 
bracts lanceolate-linear, subobtuse, 2-2.5 cm long and 4-5 mm wide. 
Ligulate florets yellow, pinkish on outer side. Achenes 7-8 mm long, 
slightly curved, ribbed, somewhat densely tuberculate-toothed along ribs; 
pappus snow-white, soft, with many bristles, 12-14 mm long, plumose, 
inner bristles barbed above. Flowering June. 

Stony-rubbly slopes. — Soviet Central Asia: Pamiro-Alai. General 
disribution: Afghanistan. Described from Fergana. Type in Leningrad. 

Series 28. Ovatae Lipsch. in Addenda XXVIII, 722. — Capitula broadly 
cylindrical. Ligulate florets yellow or pink (violet when dry). Achenes 
flattened-cylindrical, angular. Basal and lower cauline leaves ovate or 
oblong-lanceolate, with undulate-crimped or flat margin. 

95 62. S. ovata Trautv. in Tr. Peterb. Bot. Sada, I, 2 (1872) 275; Lipsch. 
Fragm. Monogr. Scorzonera I (1935) 56; Popov in Tr. Uzb. Gos. Univ. 



95 

Nov. Ser. 27, Biol. 14 (1941) 91; Lipsch. in Fl. Turkm. VII (1960) 306; 
kovalevskaja in Fl. Uzb. VI, 443 (excl. syn. 5. crassifolia Krasch. and 
Lipsch.).— Ic: Lipsch. op. cit. (1935) Plate 21 and 22; Lipsch. op. cit. 
(1960) Plate XLVII, 2. 

Perennial. Plant (9)10-2-(35) cm high. Root thickened into oblong or 
roundish tuber; root collar covered with brown or stramineous shining 
sheaths. Stems leafy, especially in lower part, branched, flowering branches 
(2)5-15(20) cm long, slightly curved, (2)3-7(15). Basal and cauhne leaves 
hard with pinnate venation, margin cartilaginous, undulate-crimped; 
lowermost leaves broadly ovate, acuminate, less often subobtuse, narrowed 
into petiole; middle cauline and those on branches reduced above, oblong- 
lanceolae, roundish at base, amplexicaul. Capitula broadly cyUndrical, 2.5- 
3.5 cm long and 0.7-1 cm wide. Involucral bracts glabrous, coriaceous, 
membranous along margin, often bordered with red stripe; outer bracts 
ovate-deltoid, obtuse; inner ones oblong-lanceolate, subobtuse, much 
longer than outer. Ligulate florets yellow, far exserted from involucre. 
Achenes 6-7 mm long, weakly curved; flattened-cylindrical, angular- 
ribbed, transversely tuberculate-serrate; pappus snow-white, dense, soft, 
its bristles plumose, barbed above. Flowering April to May. 

Clayey, sandy and stony-rubbly steppes. — Soviet Central Asia: 
mountainous Turkmenia, ? Amu-Darya, Pamiro-Alai (Zeravshan). General 
distribution: ?Iran (northern); Indo-Himalayan (Afghanistan). Described 
from vicinity of Ashkhabd. Type in Leningrad. 

Note. Var. major Bomm. and Sint. (in Lipsch. Fragm. Monogr. 
Scorzonera I (1935) 56; Lipsch. in Fl. Turkm. VII, 306) is recognized, 
which is characterized by taller, up to 35 cm high, sturdier stem, longer 
flowering shoots (10)15-25 cm long), large (up to 15 cm long) leaves, 
and larger capitula (up to 3.5 cm long and 1.5 cm wide). 

63. S. crassifolia Krasch. and Lipsch. in Lipsch. Fragm. Monogr. 
Scorzonera I (1935) 60. — 5. ovata ssp. crassifolia Krasch. and Lipsch. in 
sched., olim. — Ic: Lipsch. op. cit. (1935) Plate 23. 

Perennial. Root thickened into oblong or roundish tuber; root collar 
covered with numerous brown or stramineous, shining sheaths. Stem short, 
(7)8-12 cm long, branched from middle, branches two to four, 2-7 cm 
long, terminating in solitary capitula. Leaves glaucous-green, fleshy, 
slightly tomentose or with powdery bloom, glabrescent, flat along margin; 
basal and lower cauline leaves roadly ovate, long-petiolate, subobtuse or 
with small cusp; middle cauline leaves and those on branches reduced, 
lanceolate or oblong-lanceolate, sessile, semiamplexicaul. Capitula 
96 broadly cylindrical, (2)2.5(3) cm long and (0.8)1-1.2 cm wide. Involucre 
glabrous, glaucous; involucral bracts coriaceous, margin partially or 
completely violet; outer bracts ovate-deltoid, smaller, obtuse; inner bracts 



96 

longer, oblong-lanceolate, narrowed toward apex, subobtuse, sometimes 
with parallel veins. Ligulate florets yellow, exserted from involucre. 
Achenes 6-7 mm long, flattened-cylindrical, slightly curved, angular- 
ribbed, transversely tuberculate-toothed; pappus snow-white, soft, dpnse, 
its bristles plumose, inner bristles longer, barbed above. Flowering April 
to May. 

On conglomerates, stony slopes. — Soviet Central Asia: Pamiro-Alai 
(Fergana). Endemic. Described from Fergana. Type in Leningrad. 

Note. The species is quite close to S. ovata Trautv., from which it 
differs in having fewer flowering shoots and more fleshy, subobtuse leaves 
with flat margins. 

64. S. papposa DC. Prodr. VII, 1 (1838) 119; Boiss. Fl. or. Ill, 765; 
Post, Flora of Syr., Pal. and Syn. (1896) 493; Bordz. in Zap. Kiev. Obshch. 
Estestv. XXV, 1 (1915) 130; Bouloumoy, Fl. du Liban (1930) texte, 209; 
Grossh. Fl. Kavk. IV (1934) 238; Lipsch. Fragm. Monogr. Scorzonera I 
(1935) 64; II (1939) 17; Isaev in Fl. Azerb. VIII, 524.-5. papposa DC. 
var.? Stschegleew in Bull. Soc. Nat. Mosc. XXIV, 4 (1851)'473; XXVI, 
2 (1853) 321.— 5. paradoxa Fisch. and Mey. in DC. Prodr. VII, 1 (1838) 
119. — S. kurdica Bois. and Noe in Boiss. Diagn. pi. or. ser. II, 3 (1856) 
93. — S. picridioides Boiss. Fl. or. Ill (1875) 763 (p. min. p. quod plant. 
Szovitzian.).— Ic: Lipsch. op. cit. I (1935) Plate, 25; II (1939) Plate 1. 

Perennial. Plant weakly tomentose, glabrescent. Root thickened into 
tuber; root collar almost glabrous. Stems (8)10-18(30) cm high, mostly 
thick, erect or slightly flexuous, sulcate, more or less strongly branched 
in upper part or from base; branches (floral) simple or in turn branched, 
usually many (2)3-7(12). Basal and lower cauline leaves petiolate, varying 
in size, ovate or oblong, somewhat obtuse or acuminate; middle leaves 
sessile, semiamplexicaul, long-acuminate; uppermost leaves reduced, 
lanceolate; all leaves coriaceous, with cartilaginous margin, usually highly 
undulate-crimped. Capitula large, on long peduncles, broadly cylindrical, 
(2.5)3-^ cm long, and (0.7)1-1.5 cm wide. Involucre glabrous; bracts 
coriaceous with membranous margin, sometimes bordered with lilac- 
colored stripe, obtuse; outer bracts short-ovate or ovate; inner bracts 
oblong-lanceolate, two to two and one-half times as long as outer. Ligulate 
florets violet, greatly exserted from involucre. Achenes (7)8-10 mm long, 
sharply tetraquetrous, white, strongly tuberculate-toothed; pappus dense, 
snow-white, its bristles plumose, inner bristles barbed above. Flowering 
May to June. 
97 Dry slopes. — Caucasus: Southern Transcaucasia (Armenia, 

Nakhichevan). General distribution: Eastern Mediterranean, Armenia and 
Kurdistan, Iran. Described from Syria. Type in Geneva. 

Section 14. Hissaricae Lipsch. Fragm. Monogr. Scorzonera II (1939) 



97 

63, pro subsect. — Root thickened into tuber. Stems simple, scape-like, 
weakly leafy. Capitula cylindrical or turbinate. Involucre of 8-12 bracts. 
Achenes large and glabrous, with longitudinal scabrous ribs. Ligulate 
florets yellow; pappus bristles stiff, wholly or at base blackish violet, 
plumose throughout. 

65. S. hissarica Winkl. in Tr. Peterb. Bot. Sada XI (1889) 172; 
Lipsch. Sist. Zam. ob Aziat. Skorts. (1932) 13; Lipsch. Fragm. Monogr. 
Scorzonera II (1939) 63; Popov in Tr. Uzb. Gos. Univ. Nov. Ser. 27, 
Biol. 14 (1941) 92; Lipsch. in Fl. Turkm. VII (1960) 306; Kovalevskaja 
in Fl. Uzb. VI, 444.-5. hissariae Ind. Kew. Suppl. I (1901-1906) 388 
(lapsu calami). — Ic: Lipsch. op. cit. (1932) 31, Fig. 7; Lipsch. op. cit. 
(1939) Plate 9. 

Perennial. Root thickened into tuber; root collar covered with brown 
remnants of sheaths of basal leaves, sometimes producing two or three 
(five) stems. Stems sparsely leafy, scapelike, (6)10-20(30) cm long, 
simple, weakly pubescent, terminating in solitary capitula. Basal leaves 
broadly lanceolate or ovate-lanceolate, acuminate, slightly cartilaginous 
along margin, toothed, flat or undulate-crimped, glaucous, bloom 
subsequently disappearing, narrowed into petiole broadening at base into 
sheath; cauline leaves less numerous, sessile, lanceolate, narrower than 
basal ones. Capitula large, 2.5-3.5(4) cm long, broadly cylindrical or 
turbinate. Involucre of 8-12 bracts, pubescent or almost glabrous, outer 
bracts deltoid; inner ones ovate-lanceolate, two to three times as long as 
outer; all bracts subacute. Ligulate florets yellow, turning pink on drying. 
Achenes large, ribbed, toothed-scabrous along ribs; pappus bristles stiff, 
plumose, wholly or at base blackish-violet. Flowering April to June. 

Varicolored low hills, stony-rubbly slopes. — Soviet Central Asia: 
Pamiro-Alai. Endemic. Described from Hissar. Type in Leningrad. 

Note. The species is extremely variable in the size of the organs, 
especially the leaves. Some specimens in their habit resemble Scorzonera 
inconspicua Lipsch., from which they can be distinguished easily by the 
structure of the achene and the pappus, particularly by its color. 

Section 15. Nervosae Lipsch. in Lipsch. Fragm. Monogr. Scorzonera 
I (1935) \26.—Lasiospora Less, in DC. Prodr. VII, 1 (1838) 124, p. p. 
and in Boiss. Fl. or. Ill (1875) 756, p. p. — Euscorzonera DC. § 
Tomentosae Boiss. op. cit. Ill (1875) 756. — Large perennials with densely 
98 leafy stems. Pubescence very dense, shining (velvety) or less dense, dull, 
tomentose, ash-colored (not velvety); sometimes plants almost glabrous. 
Leaves of variable width, usually large, ovate-elliptical or lanceolate, 
venation curvinerved, veins numerous. Achenes glabrous or pubescent. 
Pappus bristles soft, plumose or stiff, barbed-scabrous. 



98 

Series 29. Sosnovskyanae Lipsch. in Addenda XXVIII, 722. — 
Pubescence very dense, shining (velutinous). Leaf veins hairy. Leaves 
not undulate along margin. Achenes pubescent or glabrous. 

66. S. sosnovskyi Lipsch. Fragm. Monogr. Scorzonera I (1935) 142; 
Grossh. Opred. Rast. Kavk. 509. — S. dzhawakhetica Sosn. in Grossh. Fl. 
Kavk. IV (1934) 236, descr. ross.; Sosnowvsk. in Tr. Tbil. Bot. Inst. II 
(1938) 218; Papava in Fl. Gruzii VIII, 645. — S. dzhawakhetica Sosn. var. 
viridescens Sosn. in Fl. Gruzii VIII (1952) 646, descr. georg. — S. nervosa 
auct. fl. cauc. p. p. quoad pi. Georgiae. — S. veratrifolia auct. fl. cauc. p. 
p. (non Fenzl). — Ic: Lipsch. op. cit. (1935) Plate 49. 

Perennial. Root thickened, vertical, cylindrical; root collar covered 
with indurescent remnants of leaf sheaths and petioles, whole plant 
velutinous-sericeous (after long preservation in herbarium, pubescence 
turns rusty). Stems numerous, hard, erect or slightly curved, leafy 
throughout, more or less highly branched (often dichotomous). Leaves 
flat, not undulate along margin, with many (7-10) veins, barely visible 
due to dense velvety pubescence; basal leaves ovate, obtuse or subobtuse, 
narrowed at base into sulcate, usually short, and slightly flattened petiole; 
lower cauline leaves with short flat petiole, slightly amplexicaul, broadly 
lanceolate or ovate, subobtuse; middle cauline leaves sessile or almost 
so, semiamplexicaul, ovate-lanceolate or lanceolate, all leaves reduced 
toward stem apex, subobtuse. Capitula turbinate, 1.8 cm long, solitary, 
on sulcate, pubescent or sometimes almost glabrous leafy branches; leaves 
often crowded near capitula. Involucre sparsely pubescent or almost 
glabrous; bracts sometimes with carinate midrib, membranous along 
margin; outer bracts small, ovate-deltoid, acuminate; inner bracts two to 
three times as long, lanceolate or oblong-lanceolate, acuminate. Ligulate 
florets yellow, slightly longer than involucre. Achenes densely white lanate; 
pappus two to two and one-half times as long as achenes, of rusty or 
yellowish hairs, plumose, barbed-scabrous above. Flowering August. 

Stony-rubbly slopes and peaks, rock crevices. — Caucasus: Eastern 
Transcaucasia (Dzhavakhetia). Endemic. Described from Akhalkalaki 
Region (Tetrobo-Chobaret Range). Type in Moscow. 

67. S. ketzkhovelii Sosn. in Lipsch. Fragm. Monogr, Scorzonera I 
(1935) 145 (descr. latin.); Grossh. H. Kavk. IV (1934) 240 (descr. rossica); 

99 Sosn. in Tr. Tbil. Bot. Inst. II (1938) 219 (descr. lain.); Grossh. Opred. 
Rast. Kavk. 510; Papava in Fl. Gruzii VIII, 647.— Ic: Fl. Gruzii VIII, 
Plate 434. 

Perennial. Very similar to S. sosnovskyi Lipsch. Root strong, vertical, 
cylindrical; whole plant very densely velutinous-sericeous. Stems 
numerous, erect, sulcate, leafy up to inflorescence, apically with spreading 
one-headed branches. Basal leaves long-petiolate, ovate-lanceolate, 



99 

obtuse, with many (up to 10) veins, entire, not undulate; lower cauline 
leaves oblong-lanceolate, short-petiolate; higher leaves sessile, lanceolate, 
subacute, gradually reduced toward apex of stem. Capitula rather large 
(up to 2 cm long), peduncles almost glabrous, greenish. Involucre almost 
glabrous; outer bracts small, ovate-lanceolate, acuminate, almost 
membranous along margin. Ligulate florets yellow. Achenes (even when 
young) completely glabrous, obpyramidal, tetraquetrous; pappus reddish- 
rusty or dirty yellow, its bristles plumose almost to apex. Flowering July. 

Stony-gravelly slopes and peaks, rock crevices — Caucasus: Eastern 
Transcaucasia (Dzhavakhetia). Endemic. Described from Akhalkalaki 
Region (Tetrobo-Chobaret Range). Type in TbiUsi. 

Note. Very close to S. sosnovskyi, from which it differs by having 
completely glabrous achenes and ovaries, analogous to what is observed 
in the species pair 5. filifolia-S. lipskyi. 

Series 30. Latifoliae Lipsch. in Addenda XXVIII, 722. — Pubescence 
less dense, dull, tomentose, ash-colored (not velvety), sometimes plants 
almost glabrous. Leaves with very prominent veins and cartilaginous, 
undulate-crimped margins. Capitula many. Inflorescence paniculate. 
Achenes pubescent or glabrous (in species growing in Asia Minor). 

68. S. latifolia (Fisch. and Mey.) DC. Prodr. VII, 1 (1838) 124; 
Grossh. Fl. Kavk. IV (1934) 236; Lipsch. Fragm. Monogr. Scorzonera I 
(1935) 133; II (1939) 24; Lipsch. in Tr. Bot. Inst. Akad. Nauk SSSR, 
Ser. 1, II (1936) 305; Grossh. Opred. Rast. Kavk. 509; Papava in Fl. 
Gruzii VIII, 646; Isaev in Fl. Azerb. VIII, 526. — Lasiospora latifolia Fisch. 
and Mey. Ind. Sem. Hort. Petrop. I (1835) 30.— 5. nervosa Trev. Delect. 
Sem. Hort. Bonnens. (1833) non Lam. nee Pers.; opusc. non vidi; Linnaea 
(1835) 73; DC. op. cit. (1838) 125; Boiss. Fl. or. Ill, 779; Medved. in Tr. 
Tifl. Bot. Sada XVIII, 2 (1918) 307; Grossh. in Bot. Shorn. Azerb. GNU, 
I (1932) 50; Prilipko in Tr. Bot. Inst. Baku II (1936) 46; Sosn. in Tr. 
Tbil. Bot. Inst. II (1938) 224. — Lasiospora nervosa Sch. Bip. in Nov. 
Acta Acad. Caes. Leopol.-Carol. Nat. Cur. XXI (1845) 166. — S. paniculata 
Boiss. Diagn. pi. or., ser. I, 7 (1846) 8. — S. lasiospora Boiss. and Balansa 
in Boiss. Diagn. pi. or. ser. II, 6 (1859) 118.— Ic: Lipsch. op. cit. (1935) 
Plates 45, 45a; Photos 6, 7; Fl. Azerb. VIII, Plate LIV. 
100 Perennial. Root vertical, strongly thickened; root collar covered with 

indurescent sheaths and remnants of previous year's stem bases; whole 
plant pubescent, gray, less often almost glabrous to glacucous-green or 
green. Stems 20 cm to 1 m high, numerous (to 20 from single root), 
strong, erect, glabrous or slightly pubescent, sulcate, becoming glabrous 
and distinctly angular toward apex, densely leafy, with many branches 
above. Leaves hard, many- veined (5-15), variable in shape and width, 
with cartilaginous margins, more or less toothed-serrate, scabrous. 



100 

undulate-crimped; basal leaves narrowed into more or less long, sulcate 
petiole, lanceolate or broadly lanceolate, attenuate-acuminate, cusp callose, 
sometimes bent; cauline leaves lanceolate, oblong, broadly ovate- 
lanceolate; lower cauline leaves short-petiolate, petioles slightly flattened 
at base; middle and upper cauline leaves sessile, amplexicaul; all leaves 
gradually reduced upward. Capitula turbinate or cylindrical, 1.8-2.3 cm 
long (at flowering) and 1.8-2.5 cm (at fruiting), numerous, solitary (rarely 
in twos or threes), terminal on lateral branches bearing few small, linear 
or subulate leaves; flowering shoots forming pyramidal-paniculate 
compound inflorescence. Involucre pubescent, bracts hard, somewhat 
carinate; outer bracts small, deltoid, acuminate, cusps often bent; inner 
bracts lanceolate, much longer than outer, acuminate. Ligulate florets 
yellow, slightly longer than capitula. Achenes terete, narrowed at base, 
7-10 mm long, densely white lanate; pappus rusty, as long as achene, its 
bristles densely plumose. Flowering June to August. 

Dry stony slopes and screes. — Caucasus: Eastern and Southern 
Transcaucasia. General distribution: Armenia and Kurdistan. Described 
from Azerbaidzhan. Type in Leningrad. 

Note. The species varies greatly in pubescence and form, as well as 
width of leaves. On the basis of pubescence, it is possible to distinguish: 
var. glabrata Trautv. {Tr. Peterb. Bot. Sada II (1873) 559)— stems in the 
upper part, upper leaves, and the involucre almost glabrous; var. pubescens 
Lipsch. nom. nov. (= var. typica Trautv. op. cit. 559) — whole plant and 
involucre more or less tomentose. Based on the form of the leaves, Prilipko 
(cf.: Lipsch. Fragm. Monogr. Scorzonera I (1935) 136 and Tr. Bot. Inst. 
Baku II (1936) 46) recognized the following varieties: var. macrophylla 
Pril. — with broad cauline oval or ovate leaves (4-5.5 cm wide); var. 
angustifolia Pril. — leaves lanceolate, 1-1.5 cm wide; var. media Pril. — 
leaves usually undulate-crimped along the margin, 2.3-5 cm wide; var. 
abbreviata Pril. — leaves narrowly lanceolate (0.8-1.2 cm wide), flowering 
branches reduced; capitula densely aggregated, few (three to nine). Many 
plants do not fit this subdivision and are intermediate in character. 

69. S. pulchra Lom. in Tr. Tifl. bot. Sada II (1897) 289; Grossh. in 

Bot. Shorn. Azerb. GNII I (1932) 50; Grossh. Fl. Kavk. IV (1934) 237; 

Lipsch. Fragm. Monogr. Scorzonera I (1935) 142; Grossh. Opred. Rast. 

101 Kavk. 509; Isaev in Fl. Azerb. VIII, 527.— Ic: Lipsch. op. cit. (1935) 

Plate 48. 

Perennial. Plant densely white tomentose. Root cyUndrical, thick; root 
collar covered with brown remnants of leaf sheaths. Stems densely white- 
tomentose, tall, (30)35-60 cm high, densely leafy, branched; branches 
ascending, slightly flexuous, varying in length, simple, axillary; forming 
almost corymbose compound inflorescence. Leaves white-tomentose, 
many-veined, veins covered by hairs, with cartilaginous border along 



101 

margin, usually undulate-crimped; basal leaves lanceolate, narrowed into 
petiole, broadened into sheath; cauline leaves sessile, weakly amplexicaul, 
broadly lanceolate or ovate, acuminate, cusps often subulate, sometimes 
curved; all leaves gradually reduced upward, uppermost leaves close to 
capitula lanceolate. Capitula (3)4-10, solitary terminal on flowering 
shoots in corymbose inflorescence, turbinate or broadly cylindrical, 
(1.5)2-2.2 cm long and 1-1.2(1.5) cm wide. Involucre white-tomentose, 
sometimes glabrescent; outer bracts smaller, deltoid-lanceolate; inner 
bracts oblong-lanceolate,, often with carinate midrib, subobtusely 
acuminate much longer than outer bracts. Ligulate florets sulfur-yellow, 
longer than involucre. Achenes 7-8 mm long, with dense woolly pube- 
scence; pappus dirty-yellow, its bristles plumose, hairs barbed above. 
Flowering June to August. 

Dry slopes. — Caucasus: Eastern Transcaucasia. Endemic. Described 
from Karabakh (Ziarat Mountain). Type in Leningrad; isotype in Tbilisi. 

Note. From the closest species S. latifolia (Fisch. and Mey.) DC, it 
is distinguished by having white-tomentose pubescence and a corymbose 
inflorescence. 

From S. sosnovskyi Lipsch. it is distinguished by having white- 
tomentose, less dense, and nonvelvety pubescence, acuminate and not 
obtuse leaves, etc. 

70. S. safievii Grossh. in Bot. Shorn. Azerb. GNU I (1932) 51, 
descr. ross.; Tr. Azerb. Otd. Zakavk. Fil. Akad. Nauk SSSR, Sekt. Bot. 
I (1932) 62, descr. latina; Grossh. Fl. Kavk. IV (1934) 237; Lipsch 
Fragm. Monogr. Scorzonera I (1935) 138 Grossh. Opred. Rast. Kavk. 
509.— Ic: Lipsch. op. cit. (1935) Plate 47. 

Perennial. Plant canescent. Root vertical; root collar covered with 
remnants of indurescent sheaths of basal leaves; sheaths often hairy on 
inner side; latex yellow. Stems 35-50 cm high, more or less pubescent, 
especially in lower part, usually many (up to eight), ascending at base, 
straight, relatively slender, sulcate, highly branched, densely leafy. Leaves 
hard, glaucous-green, linear-lanceolate, narrow, 3-5(8) mm wide, flat 
with many (three to seven) veins and subulate tip; basal leaves narrowed 
into sulcate, sometimes weakly carinate petiole; cauline leaves sessile, 
slightly amplexicaul; all leaves reduced upward. Capitula 1.5-2(2.5) cm 
102 long (at fruiting), usually many (3-20), solitary terminal on long flexuous 
branches — axillary peduncles. Involucral bracts rigid, pubescent, 
glabrescent, somewhat carinate, acuminate, with narrowly membranous 
margin; outer bracts small, deltoid, acuminate, often with bent cusps; 
inner bracts oblong-lanceolate, much longer than outer. Ligulate florets 
yellow, longer than involucre. Achenes terete, (6)7-10 mm long, densely 
white-lanate; pappus dirty yellow, its bristles rusty, densely plumose, 
barbed above. Flowering July to August. 



102 

Dry stony slope. — Caucasus: Southern Transcaucasia. Endemic. 
Described from Armenia (Daralagyoz). Type in Baku; isotype in Moscow. 

Section 16. Vierhapperia Lipsch. Fragm. Monogr. Scorzonera II 
(1939) 88. — Plants covered with lanate pubescence. Stems leafy. Leaves 
(in species growing in the USSR) usually with prominent veins. Achenes 
in our species lanate (subsect. Neovierhapperia Lipsch. nom n. = 
Euvierhapperia Lipsch.). Pappus bristles wholly plumose. 

Series 3 1 . Biebersteinianae Lipsch. in Addenda XXVIII, 722. — Stems 
ascending or erect, not virgate. Leaves hard, mostly with prominent 
veins, apex narrowed into cusp. Capitula many-flowered (more than 20); 
achenes 5-7 mm long (excluding pappus). 

71. S. ensifolia M.B. Fl. taur.-cauc. II (1808) 235 and III (1819) 
523; Ldb. Fl. Alt. IV, 162; DC. Prodr. VII, 1, 124; Ldb. Fl. Ross. II, 2, 
797; Boiss. Fl. or. Ill, 780; Schmalh. Fl. II, 144; Krasch. in Fl. Yugo- 
Vost. VI, 452; Lipsch. Fragm. Monogr. Scorzonera II (1939) 94; Kryl. 
Fl. Zap. Sib. XI, 2986. — 5. ensifolia M.B. a. glabrescens and p. pilosior 
O. Ktze. in Tr. Peterb. Bot. Sada, X, 1 (1887) 205.— 5. ensifolia var. 
lanata Serg. in Kryl. Fl. Zap. Sib. XI (1949) 2987. — Lasiospermum 
ensifolium (M.B.) Fisch. Catal. Hort. Gorenk. (1812) 34. — Lasiospora 
ensifolia (M.B.) Cass, in Diet. sc. nat. XXV (1822) 307; Sch. Bip. in 
Nov. Acta Acad. Caes. Leopol.-Carol. Nat. Cur. XXI (1845) 166.— L. 
ensifolia (M.B.) Claus in Gobel, Reise II (1838) 284. — L. macrophylla 
Fisch. and Mey. Enum. pi. nov. I (1841) 39. — Scorzonera macrophylla 
Fisch. and Mey. Enum. pi. nov. I (1841) 39 in adnot.; Ldb. Fl. Ross. II, 
2, 797; Lipsch. op. cit. (1939) 47.— Exs.: GRF No. 1167. 

Perennial. Root vertical, cylindrical; root collar sometimes covered 
with indurescent sheaths of basal leaves, densely velutinous from long 
rusty hairs. Stems 20-55 cm high, few, less often solitary, slightly 
flexuous, weakly sulcate, pubescent (often floccose in leaf axils), less 
often glabrous, densely leafy throughout, usually branched, less often 
simple. Leaves linear-lanceolate or linear, 6-25 cm long and 2-6(10) 
mm wide, apex attenuate into subulate bent cusps, hard, with many 
103 veins (rather distinct). Capitula solitary or two to four, terminal. Involucre 
more or less lanate-velutinous (sometimes pubescence forms "white 
collar" around involucre — var. pilosior O. Ktze.); involucral bracts 
lanceolate, long-acuminate; outer bracts reduced, with bent cusp. Ligulate 
florets yellow, one and one-half times as long as involucre. Achenes 
lanate; pappus dirty yellow or rusty, its bristles plumose from base, 
two to two and one-half times as long as achenes. Flowering (May) 
June to August. 

Sandy steppes, hummocky and hilly sands, dunes, sandy steppe pine 
forests, lowlands. — European Part: Volga-Don, Trans-Volga, Black Sea 



103 

Region, Lower Don, Lower Volga; Western Siberia: Upper Tobol, Irtysh, 
Altai; Soviet Central Asia: Aralo-Caspian, Balkhash Region, Kyzyl-Kum, 
Amu-Darya, Syr-Darya. General distribution: ?Dzhungaria-Kashgaria. 
Described from Caspian desert. Type in Leningrad. 

Note. Scorzonera macrophylla Fisch. and Mey. (type preserved in 
Leningrad), described from sands of the Irtysh River, is the broad-leaved 
variant of S. ensifolia M.B. 

72. S. biebersteinii Lipsch. Fragm. Monogr. Scorzonera II (1939) 
95; Isaev in Fl. Azerb. VIII, 527. — S. eriosperma M.B. Beschr. Land, 
zw. Terek und Kur. (1800) 117 and 28, non Gouan, 1773; Willd. Sp. 
pi. Ill, 3 (1803) 1504; M.B. Fl. taur.-cauc. II (1808) 236 and III (1819) 
523; DC. Prodr. VII, 1, 124; Ldb. Fl. Ross. II, 2, 796; Boiss. Fl. or. Ill, 
780; Grossh. Fl. Kavk. IV (1934) 237; Lipsch. op. cit. (1939) 95; Grossh. 
Opred. Rast. Kavk. 509; Papava in Fl. Gruzii VIII, 647. — Lasiospora 
eriosperma XM..B.) Sch. Bip. in Nov. Acta Acad. Caes. Leopol.-Carol. 
Nat. Cur., XXI (1845) 165. — Lasiospermum angustifolium Fisch. Catal. 
Hort. Gorenk. (1812) 34, nomen nudum. — Lasiospora angustifolia Cass, 
in Diet. sc. nat. XXV (1822) 306.— Ic: M.B. Cent. pi. II, dec. II (1843) 
t. 69; Fl. Azerb. VIII, Plate LIII, 2.— Exs.: Herb. R. Cauc. No. 299; PI. 
or. exs. No. 50. 

Perennial. Plant more or less lanate (woolly pubescence most 
abundant near stem base and in leaf axils); root collars sparsely covered 
with membranous sheaths and woolly pubescence. Stems 10-30(50) cm 
high, several arising from single root, ascending at base, more or less 
branched, less often simple. Leaves narrowly linear, (1)2-2.5(3) mm 
wide, with subulate apex, undulate, three(five)-veined; cauline leaves 
slightly amplexicaul. Capitula turbinate or ovoid, 1-10 (up to 22!) on 
single plant (depending upon nature of stem branching), small, 1.2- 
1.5(1.9) cm long. Involucre weakly pubescent, often almost glabrous; 
outer bracts small, deltoid, with short cusp; inner bracts lanceolate, much 
longer than outer, sometimes with distinct carinate midrib. Ligulate florets 
yellow, turning pink on drying, a quarter longer than involucre. Achenes 
obovoid, at base acute, lanate; pappus dirty yellow or rusty, its bristles 
plumose throughout. Flowering May to July. (Plate VI, Fig. 2.) 
104 Stony and clayey slopes, sands. — Caucasus: Ciscaucasia, Dagestan, 

Eastern and Southern Transcaucasia. Endemic. Described from 
Konstantinogorsk. Type in Leningrad. 

Note. The following variants are distinguished: f. lanata Lipsky — 
stems, peduncles, and involucre lanate; var. apscheronica Lipsch., highly 
branched, with relatively small capitula. There are specimens in the 
herbarium of the Botanical Institute, Academy of Sciences of the USSR, 
with the label "Turcomania, Karelin 1838;" the plants were undoubtedly 
collected from the Caucasus. 



104 

Series 32. Ramosissimae Lipsch. in Addenda XXVIII, 723. — Stems 
virgate, flexuous, erect. Leaves softer, without distinct cusp at apex. 
Capitula few-flowered (with six or seven flowers). Achenes 9-10 mm 
long (excluding pappus). 

73. S. ramosissima DC. Prodr. VII, 1(1838) 125; Boiss. and Buhse 
in Mem. Soc. Nat. Mosc. XII (1860) 139; Boiss. Fl. or. Ill, 781; Grossh. 
in Bot. Shorn. Azerb. GNII, I (1932) 51; Grossh. Fl. Kavk. IV (1934) 
237; Lipsch. Fragm. Monogr. Scorzonera II, 139; Grossh. Opred. Rast. 
Kavk. 509; Isaev in Fl. Azerb. VIII, 528. — 5. eriosperma auct. non 
M.B.: Auch. ex DC. Prodr. VII, 1 (1838) 125. 

Perennial. Plant with appressed pubescence. Root collar covered with 
indurescent, undivided or fibrous, sheaths of basal leaves and rusty woolly 
pubescence. Stems few, 25—45 cm high, straight, upright, virgate, sulcate, 
pubescent, simple below, dichotomously branched from middle. Leaves 
linear, herbaceous, with silky-rusty hairs, flat or sligjjtly folded 
lengthwise, with three indistinct veins, somewhat obtuse; basal leaves 
6-8 cm long and 1.5-2 mm wide, broadened at base into membranous, 
whitish, deltoid-lanceolate sheaths; cauline leaves reduced, filiform. 
Capitula solitary, on long slender flowering shoots, with few (six or 
seven) flowers, turbinate, 1-1.5(1.8) cm long (at flowering), 1.8-2.2 cm 
(at fruiting). Involucre pubescent or almost glabrous (var. glabrescens 
Grossh.); outer bracts small, acuminate, with blackish, slightly bent tip; 
inner bracts lanceolate, considerably longer than outer ones, with more 
or less distinct midrib. Ligulate florets yellow, slightly longer than 
involucre. Achenes 9-10 mm long, densely lanate, narrowly-terete, 
narrowed at base, ribbed, almost as long as pappus; pappus of plumose 
bristles. Flowering May to July. 

Limestone deposits. — Caucasus: Southern Transcaucasia 
(Nakhichevan ASSR). General distribution: Iran. Described from Elvend 
Mountain. Type in Geneva. 

Section 17. Tuberosae Lipsch. Fragm. Monogr. Scorzonera II (1939) 
39. — Sect. Lasiospora Less. Synops. Compos. (1832) 134, p. p.; DC. 
Prodr. VII, 1 (1838) 124, p. p.; Ldb. Fl. Ross. II, 2 (1845-1846) 796, p. 
p.; Sch. Bip. in Nova Acta Acad. Caes. Leopol.-Carol. Nat. Cur. XXI 
105 (1845) 166, p. p.; Boiss. Fl. or. Ill (1875) 776, p. p.— Plants usually 
small; root thickened into ovoid-spherical tuber. Achenes lanate. 
Receptacle pubescent (in all species?). 

Subsection 1. Lanatae Lipsch. in Addenda XXVIII, 723. — Stems 
simple, unbranched. 



105 

74. S. lanata (L.) Hoffm. in Comm. Soc. Phys.-Med. Mosq. I (1806) 
9; M.B. Fl. taur.-cauc. II (1808) 237 and III (1819) 524; DC. Prodr. 
VII, 1, 124, p. max. p.; Ldb. Fl. Ross. II, 2, 797; Boiss. Fl. or. Ill, 776, 
p. min. p.; Grossh. in Hot. Sbom. Azerb. GNU, I (1932) 50; Grossh. Fl. 
Kavk. IV (1934) 235; Lipsch. Fragm. Monogr. Scorzonera I (1939) 41; 
Grossh. Opred. Rast. Kavk. 509; Papava in Fl. Gruzii, VIII, 645; Isaev 
in Fl. Azerb. VIII, 528, p. p. — Leontodon lanatus L. Amoenit. acad. IV 
(1759) 287, excl. syn. nonn. — Tragopogon lanatus (L.) Willd. Sp. pi. 
Ill, 3 (1803) 1495.— Troximon lanatum (L.) Pers. Synops. II (1807) 
360. — Lasiospora lanata (L.) Fisch. in Bull. Soc. Nat. Mosc. VI (1833) 
257; C. Koch in Linnaea XVII (1843) 275.— L. lanata (L.) Fisch. and 
Mey. ex Lipsch. op. cit. (1939) 41, p. p. — ?L. tuberosa Cass. p. latifolia 
Sch. Bip. in Nova Acta Acad. Caes. Leopol.-Carol. Nat. Cur. XXI (1845) 
166. 

Perennial. Root thickened into ovoid-spherical tuber. Whole plant, 
especially leaves and stems, densely lanate. Stems scape-like, (7)10-22 
cm high, solitary, one-headed, erect, usually much longer than leaves. 
Leaves linear-lanceolate, margin flat or undulate-crimped; basal leaves 
sheathed at base, often half as long as stem, with indistinct veins, mostly 
lanate (sometimes veins rather prominent on clearing woolly pubescence). 
Capitula large, up to 2-2.5 cm long (including ligulate florets). Involucre 
lanate, especially outer bracts; involucral bracts lanceolate, outer ones 
almost half or two-thirds as long as inner ones. Ligulate florets one and 
one-half to two times as long as involucral bracts, lemon-yellow. Achenes 
narrowed at base, lanate; pappus rusty, its bristles stiff, plumose, barbed 
above; receptacle pubescent. Flowering May to June. 

Dry slopes. — Caucasus: Dagestan. Endemic. Described from Derbent. 
Type in Moscow. 

75. S. pseudolanata Grossh. in Tr. Bot. Inst. Baku II (1936) 258, 
descr. latin.; Grossh. Fl. Kavk. IV (1934) 235, deer, ross.; Lipsch. Fragm. 
Monogr. Scorzonera II (1939) 42; Grossh. Opred. Rast. Kavk. 509.— 5. 
lanata auct. non Hoffm.; DC. Prodr. VII, 1 (1838) 124, p. min. p.; 
Boiss. Fl. or. Ill (1875) 776, quoad pi. Iran.; Isaev in Fl. Azerb. VIII, 
528, p. p. — S. lanata ssp. pseudolanata (Grossh.) Lipsch. op. cit. (1939) 
43, in syn. 

Perennial. Root tuberous. Whole plant, especially leaves, lanate. 
Stems scape-like, short, 3-6(10) cm high, at fruiting bent, drooping, 
106 often procumbent. Leaves linear-lanceolate, up to 0.6-0.8 cm wide, as 
long as or slightly shorter than stems, flat along margin, sometimes 
more or less undulate-crimped, often uncinate. Capitula 0.4-1(1.2) cm 
long (at flowering). Involucre with lanceolate, slightly pubescent bracts; 



106 

outer ones (particularly at flowering) densely lanate, smaller; inner ones 
less pubescent, longer than outer. Ligulate florets orange-yellow, barely 
exserted from involucre (always ?). Achenes lanate; pappus rusty, its 
bristles plumose. Flowering April to June. 

Dry slopes. — Caucasus: Southern Transcaucasia. General 
distribution: Armenia and Kurdistan, Iran. Described from Armenia and 
Nakhichevan. Type in Baku. 

Note. A small race, very close to S. lanata (L.) Hoffm., with a 
series of transitional forms. Distinguished from S. lanata (L.) Hoffm. 
by its shorter size, bent, often drooping or procumbent stems, leaves as 
long as the stems, smaller capitula, and orange-yellow ligulate florets 
barely exserted from the involucre. 

Subsection 2. Tuberiferae Lipsch. in Addenda XXVIII, 723. — Stems 
branched. 

Series 33. Nutantes Lipsch. in Addenda XXVIII, 723. — Peduncles 
pendant. Leaves softer, with somewhat prominent veins. 

76. S. tuberosa Pall. Reise III (1776) 757; M.B. Fl. taur.-cauc. II, 
236; Ldb. Fl. Alt. IV, 163; DC. Prodr. VII, 1, 118; Ldb. Fl. Ross. II, 2, 
796; Boiss. Fl. or. Ill, 777, p. p.; Lipsch. and Krasch. in Byull. Mosk. 
Obshch. Isp. Prir. Otd. Biol. XLIII, 1 (1934) 139; Lipsch. Fragm. Monogr. 
Scorzonera II (1939) 48; Kryl. Fl. Zap. Sib. XI, 2985; Lipsch. in Spisok 
Rast. Gerb. Fl. SSSR XIII, 132.— 5. tuberosa var. canescens Bge. Beitr. 
z. Kenntn. Fl. Russl. u. Stepp. Centr.-As., i. e. Lehmann reliq. bot. etc. 
(1851) 200 (seorsum impressum) and in Mem. Acad. Sc. Petersb. div. 
sav. VII (1854) 376. — Lasiospora tuberosa (Pall.) Claus in Cobel, Reise 
(1838) 284. — L. tuberosa a. angustifolia Sch. Bip. in Nova Acta Acad. 
Caes. Leopol.-Carol. Nat. Cur. XXI (1845) 166.— Ic: Pallas, Reise III 
(1776) t. Y, fig. 3; Lipsch. and Krasch. op. cit. (1934) 139, Fig. 1; 
Lipsch. op. cit. (1939), Plate 4, Fig. 1.— Exs.: GRF No. 4000. 

Perennial. Small spring desert plant with tuberous root; root collar 
covered with gray or rusty woolly pubescence and less numerous, shining, 
membranous scales. Stems solitary, less often 2-5, straight or slightly 
flexuous, (3)4.5-8.5(11) cm high, pubescent, usually branched, with two 
or three slender, flowering branches. Leaves usually longer than stem 
or shorter, Unear or linear-lanceolate, 1-3(5) mm wide, slightly pubescent 
or almost glabrous, with three indistinct veins. Capitula small, cylindrical 
107 or turbinate, 3-8(15) on single plant. Involucre slightly pubescent, bracts 
membranous along margins, sometims reddish-violet; outer bracts small, 
deltoid or triangular-ovate, appressed to inner bracts, mostly with uncinate 
cusps; inner bracts lanceolate. Ligulate florets yellow, reddish when dry. 



107 

slightly exserted from involucre. Achenes lanate; pappus plumose, its 
bristles sometimes violet at tips, barbed-scabrous. Flowering April to 
May. 

Steppes, sands. — European Part: Lower Volga, Trans-Volga 
(southern part); Western Siberia: Upper Tobol, Irtysh; Soviet Central 
Asia: Aralo-Caspian Region. Described from Caspian semidesert. Type 
in London. 

77. S. sericeo-lanata (Bge.) Krasch. and Lipsch. in Byull. Mosk. 
Obshch. Isp. Prir. Otd. Biol. XLIII, 1 (1934) 141; Lipsch. Fragm. Monogr. 
Scorzonfera II (1939) 49; Kovalevskaja in Fl. Uzb. VI, 444. — S. tuberosa 
var. sericeo-lanata Bge. Beitr. z. Kenntn. d. Fl. Russl. und Stepp. Centr.- 
As., i. e. Lehmann reliq. bot. etc. (1851) 200 (seorsum impressum) and 
in Mem. Acad. Sc. Petersb. div. sav. VII (19540 376. — S. tuberosa var. 
graminifolia Winkl. ex Lipsch. and Krasch. op. cit. (1934) 141, pro 
syn.. — S. tuberosa var. polycephala Rgl. and Schmalh. ex Lipsch. and 
Krasch. op. cit. (1934) 141, pro syn. — 5. tuberosa subsp. sericeo-lanata 
Krasch. and Lipsch. in Lipsch. op. cit. (1939) 49, pro syn. — Ic: Lipsch. 
and Krasch. op. cit. (1934) 142, Fig. 2; Lipsch. op. cit. (1939) Plate 4, 
Figs. 2-3. 

Perennial. Root tuberous. Stems (8.5)10-20(25) cm high, solitary 
or two to five, erect or somewhat flexuous, sulcate, lanate, (hairs long, 
white, soft, silky, and mostly divergent), branched; branches somewhat 
bent. Leaves linear, 2-5 mm wide, attenuate-acuminate, lanate, with 
three to five indistinct veins, flat along margin; cauline leaves sessile, 
sometimes slightly amplexicaul. Capitula 5-7(25) on single plant, 
turbinate or less often cylindrical, on straight or somewhat bent peduncles 
(at flowering); peduncles pendant at fruiting. Involucral bracts pubescent, 
membranous along margin; outer deltoid, one-third to half, sometimes 
one-fifth as long as inner, lanceolate, appressed to inner bracts, cusps 
mostly straight. Ligulate florets yellow, slightly exserted from involucre. 
Achenes 5-7 mm long, densely lanate; pappus 10-12(20) mm long, its 
bristles plumose, barbed-scabrous above. Flowering April to May. 

Sands. — Soviet Central Asia: Aralo-Caspian, Balkhash Region, Kara- 
Kum, Kyzyl-Kum, Amu-Darya; Western Siberia: Irtysh. General 
distribution: Dzhungaria-Kashgaria (Kuldzha), Afghanistan. Described 
from sands of Kyzyl-Kum. Type in Leningrad. 

78. S. bungei Krasch. and Lipsch. in Byull. Mosk. Obshch. Isp. 
Prir. Otd. Biol. XLIII, 1 (1934) 144; Lipsch. Fragm. Monogr. Scorzonera 
II (1939) 52; Kovalevskaja in Fl. Uzb. VI, 444. — S. tuberosa var. crispa 
Bge. Beitr. z. Kenntn. d. Fl. Russl. u. Stepp. Centr.-As. i. e. Lehmann 

108 reliq. bot. etc. (1851) 200 (seorsum impressum) and in Mem. Acad. Sc. 
Petersb. div. sav. VII (1854) 376. — S. tuberosa subsp. bungei Krasch. 



108 

and Lipsch. in Lipsch. op. cit. (1939) 52, pro syn. — Ic: Lipsch. and 
Krasch. op. cit. (1934) 145, Fig. 3; Lipsch. op. cit. (1939) Plate 5. 

Perennial. Root tuberous; root collar sometimes swollen, covered 
with woolly pubescence and numerous membranous scales, falling off 
later. Stems 5(10)- 15(1 8) cm high, solitary or two or three, sulcate, 
canescent, distinctly branched; branches axillary, terminating in solitary 
capitula. Leaves broadly lanceolate (basal) to oblong-lanceolate, (0.4) 
0.5-1(2) cm wide, flat or slightly undulate, with appressed hairs on 
both sides and indistinct veins; basal leaves narrowed into petioles 
broadened into sheaths; cauline leaves sessile, semiamplexicaul, reduced 
upward; leaves on peduncles filiform. Capitula 3-10(18), cylindrical when 
young, later turbinate; peduncles more or less bent during flowering, 
pendent on fruting. Involucre canescent; outer bracts small, one-fifth to 
one-fourth as long as inner, mostly uncinate at apex; inner ones 
lanceolate, narrowly membranous along margin. Ligulate florets light 
yellow. Achenes 6-7 mm long, whitish-lanate; pappus 9 mm long, its 
bristles plumose, hairs barbed-scabrous above. Flowering March to April 
(May). 

Rubbly-stony outcrops. — Soviet Central Asia: Kyzyl-Kum (outlier 
ranges of Bukantau, Sultan-Uiz-Dag, etc.). Endemic. Described from 
Sultan-Uiz-Dag. Type in Leningrad. 

Series 34. Erectae Lipsch. in Addenda XXVIII, 723. — Peduncles 
upright. Leaves hard with prominent veins. 

79. S. gageoides Boiss. Diagn. pi. or. ser. II, 3 (1856) 95; Boiss. 
Fl. or. Ill, 777; Lipsch. Fragm. Monogr. Scorzonera II (1939) 45; 
Kovalevskaja in Fl. Uzb. VI, 445. — 5. macrophylla auct. non Fisch. and 
Mey,: Bge. Beitr. z. Kenntn. d. Fl. Russl. u. Stepp. Centr.-As., i. e. 
Lehmann reliq. bot. etc. (1851) 200 (seorsum impressum) and in Mem. 
Acad. Sc. Petersb. div. sav. VII (1854) 376.-5. fischeriana Krasch. 
and Lipsch. in Byull. Mosk. Obshch. Isp. Prir. Otd. Biol. XLIII, 1 (1934) 
146. — S. crassinervia Fisch. ex Lipsch. and Krasch. op. cit. (1934) 146, 
nomen. — S. tuberosa var. crispa Rgl. ex Lipsch. and Krasch. op. cit. 
(1934) 146, pro syn., non Bge. — Ic: Lipsch. and Krasch. in Byull. Mosk. 
Obshch. Isp. Prir. Otd. Biol. XLIII, 1 (1934) 147, Fig. 4; Lipsch. op. 
cit. (1939) Plate 3. 

Perennial. Root cylindrical, tuberous; root collar (very rarely 
thickened) lanate and covered with less numerous membranous scales, 
later drying out. Stems solitary (less often two or three), 5-20 cm high, 
distinctly sulcate, leafy, whitish tomentose- or lanate, branched; branches 
axillary at acute angles, terminating in solitary capitula. Leaves hard, 
109 almost coriaceous, pubescent on both sides, flat, (3)5-11 veined, veins 
hard, white, very prominent, basal leaves broadly lanceolate, less often 



109 

ovate, (0.7)2-3 cm wide and (3)10-15(17.5) cm long, almost as long as 
stem or very rarely even exceeding it, narrowed at base into petiole, 
long-acuminate, often with dry cusp, sometimes spiny, weakly bent; 
cauline leaves alternate or (at point of branching) sometimes almost 
opposite, reduced upward, more or less broadly lanceolate, usually longer 
than axillary flowering shoots. Capitula usually many, (2) 3-14 (30), in 
indistinct corymbose inflorescence, erect. Involucral bracts coriaceous, 
pubescent, lanceolate, membranous along margin, their cusps more or 
less curved; inner bracts almost two times as long as outer. Ligulate 
florets yellow, longer than involucre. Achenes 7-8 mm long, densely 
lanate; pappus of plumose bristles. Flowering April to May. 

Stony-rubbly slopes, sands. — Soviet Central Asia: Kyzyl-Kum (outlier 
ranges). General distribution: Iran-Afghanistan (Baluchistan). 5. 
gageoides Boiss. described from Baluchistan. Type in Geneva. 

Note. 5. fischeriana Krasch. and Lipsch. described from Kyzyl-Kum. 
Type in Leningrad. 

80. S. circumflexa Krasch. and Lipsch. in Byull. Mosk. Obshch. 
Isp. Prir. Otd. Biol. XLIII, 1 (1934) 148; Lipsch. Fragm. Monogr. 
Scorzonera II (1939) 54; Kovalevskaja in Fl. Uzb. VI, 445. — S. tuberosa 
var. crispa auct. fl. turkest. pro max. parte, non Bge. — S. lanata auct. 
non Hoffm.: Herd, in Bull. Soc. Nat. Mosc. XLV (1872) 370.— ?5. 
tuberosa var. glaberrima Bge. Beitr. z. Kenntn. d. Fl. Russl. u. Stepp. 
Centr.-As. i. e. Lehmann reliq. bot. etc. (1851) 200 (seorsum impressum) 
and in Mem. Acad. Sc. Petersb. div. sav. VII (1854) 376. — S. tuberosa 
subsp. circumflexa Krasch. and Lipsch. in Lipsch. op. cit. (1939) 54 pro 
syn. — Ic: Lipsch and Krasch. op. cit. (1934) 150, Fig. 5; Lipsch. op. 
cit. (1939) Plate 6. 

Perennial. Root cylindrical, tuberous; root collar (sometimes 
thickened) covered with rusty or grayish-woolly pubescence and less 
numerous shining membranous scales. Stems (5)8-15(25) cm high, 
solitary or two or three, more or less sulcate, pubescent, leafy, distinctly 
branched (very rarely simple); branches with solitary capitula. Leaves 
hard, almost coriaceous, pubescent on both sides or sericeous-lanate, 
along margins with cartilaginous stripe and usually highly undulate- 
crimped, acuminate, often rolled; lamina three- to seven- veined, veins 
hard, rather prominent; basal leaves mostly broadly lanceolate, (0.5) 
0.8-1.5 cm wide, narrowed into petiole; cauUne leaves sessile, oblong- 
lanceolate, reduced upward, alternate or sometimes opposite at point of 
branching. Capitula (2)4-10 per plant, usually on long, straight peduncles, 
in indistinct corymbose inflorescence. Involucre pubescent; involucral 
bracts membranous along margin; outer ones small, inner 
110 bracts two or three times as long as outer. Achenes 6-7 mm long, densely 



no 

lanate; pappus up to 9 mm long, its bristles plumose, barbed-scabrous 
above. Flowering April to May. 

Stony-rubbly slopes of mountains, outcrops of conglomerates. — 
Soviet Central Asia: Syr-Darya, Pamiro-Alai, Tien Shan. General distri- 
bution: Afghanistan. Described from Fergana. Type in Leningrad. 

Note. In places of contact with S. tuberosa Pall, and S. sericeo- 
lanata Krasch. and Lipsch. it "loses its purity" (leaves less hard, narrower, 
veins not so prominent, and peduncles pendant); however, the main 
characteristic — undulate leaf margins — is maintained. This tendency is 
observed in the Chu-Ili Mountains, Syr-Darya Karatau and other places. 

81. S. litvinovii Krasch. and Lipsch. in Byull. Mosk. Obshch. Isp. 
Prir. Otd. Biol. XLIII, 1 (1934) 153 (5. litwinowi); Lipsch. Fragm. 
Monogr. Scorzonera II (1939) 54; Lipsch. in Fl. Turkm. VII (1960) 
309. — S. tuberosa auct. fl. turkest. quoad, plant turcom. — S. tuberosa 
ssp. litvinovii Krasch. and Lipsch. in Lipsch. op. cit. (1939) 54, pro 
syn. — S. codringtonii Rech. f. in Oesterr. Hot. Zeitschr. 97 (1950) 263; 
Symb. Afghan. II (1955) 191.— Ic: Lipsch. and Krasch. op. cit. (1934) 
154, Fig. 6; Lipsch. op. cit. (1939) Plate 7; Lipsch. op. cit. (1960) Plate 
XL VII, 1; Rech. f. op. cit. (1955) fig. 132, 137. 

Perennial. Root tuberous; root collar more or less densely covered 
with rusty-woolly pubescence and less numerous membranous scales 
that subsequently drop off. Stems one to three, erect or slightly bent, 
(7)12-25(30) cm high, pubescent, leafy, branched; branches usually 
slightly bristly, often virgate after flowering, upright, terminating in 
solitary capitula. Leaves sessile, semiamplexicaul, hard, almost 
coriaceous, linear-lanceolate, (2)3-5(10) mm wide, long-attenuate at apex, 
almost subulate, flat, less often somewhat undulate, pubescent beneath, 
less often lanate, with three to five prominent veins; all leaves reduced 
upward. Capitula (3)7-12 on single plant, cylindrical or turbinate. 
Involucre pubescent; bracts slightly membranous along margin; outer 
bracts half as long as inner ones, lanceolate, more or less appressed to 
latter. Ligulate florets yellow, turning pink on drying. Achenes 5-7 mm 
long, lanate; pappus 10-13 mm long, plumose, its bristles plumose, 
reddish, barbed-scabrous above. Flowering April to May. 

Steppes, sands. — Soviet Central Asia: mountainous Turkmenia, 
Pamiro-Alai. General distribution: Iran, Afghanistan. Described from 
vicinity of Kushka. Type in Leningrad. 

82. S. oxiana M. Pop. in Tr. Uzb. Gos. Univ. Nov. Ser. 27, Biol. 
14 (1941) 91. 

Perennial. Root thickened into irregular tuber strongly tuberculate 
in lower part. Stems 5-7 cm high. Leaves narrowly lanceolate, without 



Ill 

prominent longitudinal veins. Capitula mostly solitary. Involucre 8-10 
mm long. 
Ill Juniper forests up to subalpine zone, stony slopes of semisteppe 

zone. — Soviet Central Asia: Pamiro-Alai. Described from the Sangi- 
Dzhuman and Kuli-Kalon mountains (Kugistan). Type in Tashkent (?). 
Note. I did not see specimens of this species. Kovalevskaja (Fl. 
Uzb. VI (1962) 444) relegated this species to synonymy under S. sericeo- 
lanata Krasch. and Lipsch. Such a treatment raises doubt. Therefore, 
tentatively I am retaining this, to me, obscure species so as to attract 
the attention of collectors. 

GENUS 1636. Epilasia (Bge.) Benth.'^ 

Benth and Hook. f. Gen. pi. II (1873) 532; Bge. Beitr. z. Kenntn. 
Fl. Russl. u. Stepp. Centr.-As., i. e. Lehmann reliq. bot. etc. (1851) 
200-203 (seorsum impressum) and Mem. Acad. Sc. Petersb. div. sav. 
VII (1854) 376-379, pro sect, generis Scorzonerae; Clark, Comp. Ind. 
(1876) 279, pro gen.; Hook. f. in Fl. Brit. Ind. Ill (1881) 419, pro gen.; 
Grossh. in Bot. Shorn. Azerb. GNII I (1932) 56, pro gen.; Lipsch. Fragm. 
Monogr. Scorzonera II (1939) 27-34, pro gen.; Sojak in Nov. bot. Horti 
Bot. Univ. Frag. (1962) 49, pro gen. 

Capitula ovoid-cylindrical. Involucre two-rowed; outer bracts three 
to six, leafy, often longer than inner or almost as long, erect or divergent; 
inner bracts coriaceous-membranous, less numerous. Receptacle flat, 
glabrous. Corolla ligulate, five-toothed, pale yelow, pink or lilac when 
dry; outer ligulate florets longer, exserted from involucre; inner ones 
much smaller than outer. Anthers sagittate at base, auricoles pointed or 
short-setose. Stigmatic branches slender. Achenes black or gray, terete, 
basally slightly broadened into short hollow stalk, ribbed, sometimes 
without ribs; ribs (5-10), retrorsely barbate; achenes of two types: 1) 
with apical ring from which pappus arises; and 2) callous ring in middle 
of achene, its upper half covered with woolly hairs, continuing into 
pappus. Pappus dense, ash gray or rusty, five of its brisdes (rarely more) 
fragile, barbed at tip, rest long-plumose. Annuals, with soft grayish 
pubescence, less often glabrous. 

Type of genus: E. hemilasia (Bge.) Clarke. 
The genus includes three or four species. 



'Treatment by S.J. Lipschitz. 

^From the Greek words: epi — above; and lasios — densely woolly; from the nature of 
the upper half of achenes. 



112 

1. Outer ligulate florets large, up to 2.5 cm long, highly exserted 
from involucre; inner ligulate florets numerous, much shorter (one- 
third to half) than outer; all florets pink or lilac 

1. E. mirabilis Lipsch. 

+ Outer ligulate florets smaller, slightly exserted from involucre; inner 

112 ligulate florets numerous, shorter than peripheral ones; all florets 

pale yellow (cream-colored) 2. 

2. Callous ring at tip of achene, upper part of achene not covered 
with woolly hairs; ribs of achene usually inconspicuous (except 
lower part) 2. E. acrolasia (Bge.) Clarke 

+ Callous ring in middle of achene, upper part of achene covered 
with woolly hairs, continuing into pappus; ribs of achene usually 
rather conspicuous 3. E. hemilasia (Bge.) Clarke 

1. E. mirabilis Lipsch. in Tr. Bot. Sada Mosk. Univ. 3 (1940) 176; 
in Fl. Turkm. VII (1960) 310; Fragm. Monogr. Scorzonera II (1939) 
37, pro syn.; Kovalevskaja in Fl. Uzb. VI, 447. — E. aerolasia var. 
macrocephala Lipsch. op. cit. (1939) 37. — E. acrolacia var. ammophila 
auct. non Kuntze; Lipsch. op. cit. (1939) 37, p. p. — IScorzonera 
(Epilasia) monocarpica Kitam. in Acta Phytotax. et Geobot. XVII (1957) 
38; Ej. Fl. Afghan. (1960) 441.— Ic: Lipsch. op. cit. (1960) Plate XLVIII, 
Fig. 1; Kitam. op. cit. (1960) fig. 102. 

Annual. Stem simple or branched, erect or ascending, often solitary, 
less often several, glabrous or weakly pubescent, 7-40 cm high. Leaves 
ovate-oblong or lanceolate, more or less pubescent; basal and lower 
' cauline leaves petiolate; all leaves somewhat amplexicaul, cartilaginous 
along margin, finely toothed. Outer hgulate florets large, up to 2.5 cm 
long, considerably exserted from involucre, pink or lilac when dry; inner 
ligulate florets much smaller (one - third to half as long as outer ones). 
Involucre almost glabrous or somewhat pubescent, outer bracts leafy, as 
long as inner ones or much longer. Achene terete, with callous ring at 
tip leaving upper part of achene not covered by woolly hairs; mature 
achenes black, often weakly pubescent, especially along ribs, with short 
hollow stalk; ribs sometimes inconspicuous, in lower part always 
conspicuous, with small spinules. Flowering April to May. 

Sands, clay, gypsiferous outcrops, loess hills. — Soviet Cental Asia: 
Kara-Kum, mountainous Turkmenia, Amu-Darya, Pamiro-Alai. General 
distribution: Afghanistan, ?Iran. Described from Soviet Central Asia. 
Type in Leningrad. 

Note. I did not see the type of Scorzonera monocarpica Kitam. — 
the species described from Nuristan (Afghanistan); it is preserved in the 
herbarium of Kyoto University. Judging from the description and figure, 
S. monocarpica evidently is identical to E. mirabilis. 



113 

2. E. acrolasia (Bge.) Clarke, Comp. Ind. (1876) 279; Lipsch. 
Fragm. Monogr. Scorzonera II (1939) 35 (excl. var. macrocephala 
Lipsch.) and in Fl. Turkm. VII, 311; Kovalevskaja in Fl. Uzb. VI, 447.— 
E. acrolasia (Bge.) Sojak in Nov. bot. Horti Hot. Univ. Prag. (1962) 
113 49. — Scorzonera acrolasia Bge. Beitr. z. Kenntn. Fl. Russl. u. Stepp. 
Centr.-As., i. e. Lehmann reliq. bot. etc. (1851) 202 (seorsum impressum) 
and Mem. Acad. Sc. Petersb. div. sav. VII (1854) 378; Boiss. Fl. or. Ill, 
782. — Epilasia hemilasia var. acrolasia Kuntze in Tr. Peterb. Bot. Sada 
X, 1 (1887) 201, in adnot. — Scorzonera acrolasia var. angustifolia Winkl. 
in Tr. Peterb. Bot. Sada XI, 1 (1890) 151. — Scorzonera ammophila Bge. 
op. cit. (1851) 202 and op. cit. (1854) 378; Boiss. Fl. or. Ill, 782.— 
Epilasia ammophila (Bge.) Clarke, Comp. Ind. (1876) 279. — Epilasia 
bungeana Clarke op. cit. (1876) 279, p. p. — Epilasia acrolasia var. 
ammophila Kuntze op. cit. (1887) 201, in adnot.; Lipsch. op. cit. (1939) 
37, p. p.— Ic: Fl. Uzb. VI, Plate XLIV, Fig. 1. 

Annual. Canescent or almost glabrous [var. ammophila (Bge.) 
Kuntze] plants. Stems 6-35 cm high, solitary, or several, erect, ascending 
or less often even prostrate, simple or branched, sulcate, often hollow 
in lower part, weakly pubescent, at places floccose-tomentose, 
glabrescent. Leaves lanceolate or ovate-lanceolate, (1)2.5-5 cm long and 
0.3-2 cm wide; basal and lower cauline leaves narrowed into distinct, 
usually flattened, petiole; upper cauline leaves sessile; all leaves 
somewhat amplexicaul, with three to five prominent veins, margin flat 
or undulate-crimped, more or less small toothed (seen under hand lens!), 
obtuse or acute, cartilaginous. Capitula ovoid-cylindrical, usually 
numerous, less often solitary, terminal on stem and branches; peduncles 
slightly thickened below capitula. Involucre more or less pubescent, 
glabrescent, sometimes glabrous (var. ammophila (Bge.) Kuntze); outer 
involucral bracts leafly, linear-lanceolate, with somewhat ciliate, 
triangular or ovate base, small-toothed along margin, apex with 
cartilaginous, sometimes bent cusp, shorter than inner bracts, as long or 
distinctly longer (var. ammophila (Bge.) Kuntze); inner involucral bracts 
15-20 mm long, slighdy membranous along margin. Outer ligulate florets 
pale yellow (cream-colored), exserted from involucre; inner florets smaller 
than outer. Achenes black, cylindrical, slighdy narrowed toward base, 
glabrous or weakly pubescent, usually with indistinct ribs (sometimes 
ribs rather prominent, particularly distinct in lower and hollow part of 
achene stalk), spinulose or hairly along ribs; callous ring apical, leaving 
upper half of achene bare, not covered by pappus wool, as in Epilasia 
hemilasia; pappus gray, dense; five (sometimes more) of its bristles 
longer, fragile, barbed; rest long-plumose. Flowering April to May. 

Sands, clayey soils, gravels. — Soviet Central Asia: Kara-Kum, Amu- 
Darya, Aralo-Caspian, Balkhash Region, Syr-Darya. General distribution: 



114 

Iran, ?Afghanistan, ?Indo-Himalayas. Described from Kyzyl-Kum. Type 
in Paris; isotype in Leningrad. 

Note. E. ammophila (Bge.) Clarke was described from Yan-Darya; 
type in Paris. 

114 3. E. hemilasia (Bge.) Clarke, Comp. Ind. (1876) 279; Kuntze in 

Tr. Peterb. Hot. Sada X, 1 (1887) 202, p. p. (incl. var. multicostata 
Kuntze and f. typica Kuntze); Kipsch. Fragm. Monogr. Scorzonera II 
(1939) 31; Lipsch. in Fl. Turkm. VII, 311; Kovalevskaja in Fl. Uzb. VI, 
446; Aliev in Fl. Azerb. VIII, 530. — Scorzonera hemilasia Bge. Beitr. 
z. Kenntn. Fl. Russl. u. Stepp. Centr.-As., i. e. Lehmann reliq. bot. etc. 
(1851) 201 (seorsum impressum) and Mem. Acad. Sc. Petersb. div. sav. 
VII (1854) 377; Boiss. Fl. or. Ill, 781. — Epilasia hemilasia (Bge.) Grossh. 
in Bot. Sbom. Azerb. GNU I (1932) 56; Grossh. Fl. Kavk. IV (1934) 
242. — Scorzonera cenopleura Bge. op. cit. (1851) 202 and op. cit. (1854) 
378; Boiss. Fl. or. Ill, 782. — Epilasia cenopleura (Bge.) Clarke op. cit. 
(1876) 279. — E. hemilasia var. (vel. monstrum) cenopleura Lipsch. op. 
cit. (1939) 33. — E. cenopleura (Bge.) Sojak in Nov. bot. Horti Bot. 
Univ. Prag. (1962) 49. — Scomozera intermedia Bge. op. cit. (1854) 377- 
378. — Epilasia hemilasia var. intermedia (Bge.) Boiss. Fl. or. Ill (1875) 
782; Lipsch. op. cit. (1939) 33. — Epilasia intermedia (Bge.) Clarke op. 
cit. (1876) 219.— E. intermedia (Bge.) Sojak op. cit. (1962) 49.— Epilasia 
bungeana Clarke, op. cit. (1876) 279, p. p. — Scorzonera nana Boiss. 
and Buhse in Nouv. Mem. Soc. Nat. Mosc. XII (1860) 138; Boiss. Fl. 
or. Ill, 782. — Epilasia hemilasia var. nana (Boiss. and Buhse) Kuntze, 
op. cit. (1887) 202, in adnot.; Lipsch. op. cit. (1939) 33. — Epilasia nana 
(Boiss. and Buhse) Grossh. in Bot. Sbom. Azerb. GNU I (1932) 56; 
Grossh. Fl. Kavk. IV (1934) 242.— Ic: Lipsch. op. cit. (1939) Plate 1, 
Fig. 1-2 (Type E. cenopleura); Plate 2, Fig. 3-4 (Type E. nana). 

Annual. Plant sometimes more or less soft floccose, sometimes — 
less often — almost glabrous. Stems solitary or more frequently several 
(three to five), highly variable in height, (5)10-30(50) cm high, erect or 
ascending (sometimes prostrate), hollow (especially in lower part), 
sulcate, simple or often branched, leafy. Leaves variable in shape, finely 
toothed under hand lens, scabrous, with somewhat prominent veins; basal 
leaves linear-lanceolate or oblong-ovate, narrowed into petiole; cauline 
leaves often less numerous (and then stems with rather long intemodes), 
lanceolate or linear-lanceolate, lower leaves with fairly long petioles, 
upper leaves sessile. Capitula usually numerous, very rarely solitary, 
ovoid-cylindrical, up to 1.5 cm long at fruiting. Outer involucral bracts 
leafy, unequal, with broadened lower part, acuminate, with more or less 
divergent tips; all involucral bracts more or less pubescent or glabrous, 
sometimes much longer than capitulum, sometimes almost as long; inner 



115 

involucral bracts five, coriaceous, slightly membranous along margin, 
obtuse, almost glabrous or arachnoid - pubescent. Outer iigulate florets 
slightly exserted from capitulum, light yellow (cream-colored), inner 
smaller. Achenes slightly broadened at base into hollow stalks, with 
many whitish hard ribs, covered with retrorse making them highly 
115 scabrous; callous ring in middle of achene, very rarely slightly higher; 
upper half of achene densely covered with woolly hairs, continuing into 
pappus; shining after removal of wool, dark brown; visible lower half 
of achene 3-4 mm long; pappus dense, gray; five (sometimes more) of 
its bristles barbed above, fragile, others long-plumose. Flowering April 
to May. 

Sands, clays, weedy places. — Caucasus: Southern Transcaucasia 
(Nakhichevan — var. nana (Boiss. and Buhse) Kuntze); Soviet Central 
Asia: Kara-Kum, mountainous Turkmenia, Amu-Darya, Syr-Darya, Aralo- 
Caspian, Balkhash Region, Pamiro-Alai. General distribution: Iran, 
?Afghanistan, China (Dzhungaria). Described from Soviet Central Asia. 
Type in Leningrad. 

Note. A highly polymorphic species varying in stem height, shape 
and size of leaves. Dwarf plants 4-6 cm high, occur — this is var. nana 
(Boiss. and Buhse) Kuntze. Described as Scorzonera nana from the 
Caucasus; type in Leningrad. Sometimes very tall forms are found, 
reaching 50 cm in height with spreading stems and branches and 
relatively large capitula (f. vegeta Sint.). The pubescence of the stems 
and leaves is also quite polymorphic (if glabrous, then it is var. subglabra 
Winkl.). The sculpturing of the achene is variable (number of ribs and 
presence or absence of spinules on them); when the ribs are smooth, 
flat and slightly hollow, it is var. cenopleura (Bge.) Lipsch. The latter 
was described by Bunge from Kyzyl-Kum as a separate species — 
Scorzonera cenopleura Bge. Type in Leningrad. — The type of S. 
intermedia Bge. — a species described from Soviet Central Asia (between 
Agatma and Karagata) — is preserved in Paris. 

GENUS 1637. Tragopogon L.' ^ 

L. Sp. pi. (1753) 789. 

Capitula solitary, terminal on main stem and branches, many- 
flowered, large or fairly large, 1.5-9 cm long, considerably enlarged at 
fruiting; peduncles weakly or strongly thickened below capitula or not. 
Involucre one-rowed, of almost similar contiguous or distant bracts; 



'Treatment by A.G. Borisova. 

Trom the Greek words tragos — goat; and pogon — beard. 



116 

involucral bracts 5-14(16), linear or linear-lanceolate, sometimes with 
white border, sometimes uncinate at fruiting, usually glabrous, sometimes 
arachnoid pubescent. Receptacle glabrous, alveolate, without scales. 
All florets ligulate, five-toothed, tubular in lower part, bisexual, 
yellow, lemon-yellow to orange or lilac, pink to purple or dark violet, 
shorter, as long or exceeding involucre. Stamens sygenesious; stigma 
bifid. Achenes sessile, attenuate, fusiform, linear-terete or oblong-terete, 
116 angular, with acute or obtuse ribs, sulcate, shallow or deeply furrowed, 
5-, 10-angled, ribs usually covered with obtuse or acute scales and 
tubercles, sometimes smooth or almost so; achenes narrowed above into 
somewhat long, short or indistinctly developed beak, as long as, shorter 
or longer than body of achene, less often without beak; beak sometimes 
clavate or with an annulus at pappus base; pappus (tuft), wholly 
deciduous, of one row of plumose, unequal bristles, basally connate into 
ring as if forming a funnel; pappus bristles almost entirely covered with 
fine plumose, interwined hair, some (5-10) bristles longer and not 
plumose in upper part. Biennial and perennial plants with milky sap. 
Root cordlike, long, straight, cylindrical, vertical, often sparingly 
branched with slender roots, sometimes tuberous; perennial species with 
branched rhizome, caespitose; root collar glabrous or covered with 
remnants of preveious year's leaves or their fibers. Stem (10)25-130 cm 
high, simple or branched, erect, leafy, sulcate, glabrous or arachnoid, or 
covered with arachnoid-tomentose, floccose pubescence, often crowded 
on some plant parts at late stages of development: in leaf axils, below 
capitula, near stem base, etc. Setose pubescence always absent. Leaves 
alternate, simple, entire, sometimes with undulate or membranous margin, 
sessile, usually long, linear to lanceolate, sometimes oblong to oblong- 
ovate, usually acuminate, sometimes almost filiform, parallel-veined, with 
more or less broadened, often semiamplexicaul base, sometimes involute; 
middle and upper cauline leaves shorter than basal; lower leaves usually 
withering subsequently; sometimes leaves exceeding inflorescence. 

Several investigators (Poddubnaya-Arnoldi, Araratyan, Marion 
Ownbey) have studied the chromosomes of the genus Tragopogon. The 
basic number 2n = 12 has been reported for the species T. dubius, T. 
pratensis, T. orientalis, T. brevirostris, T. reticulatus, T. longirostris, T. 
marginifolius and T. porrifolius. The diploid number 2n = 24 has been 
reported for T. mirus (T. dubius x T. porrifolius) and T. miscellus {T. 
dubius X T. pratensis), amphidiploid species obtained through natural 
hybridization in the USA (Marion Ownbey, 1950, 1953, 1954). These 
authors have reported differences in the chromosomal structure in 
different species with the same chromosome number. 

Type of genus: Tragopogon pratensis L. 



117 

The largest number of species of this genus is distributed in the 
Mediterranean Region; a fairly large number of species grows in Soviet 
Central Asia and the Caucasus. All Tragopogon species are endemic to 
the Old World, Europe and Asia. They were introduced into America. 
The genus comprises about 150 species. 
117 Economic Importance. The stem, leaves and roots of some species 

(e.g., T. pratensis, T. dubius, etc.) are edible and suitable for making 
salad. T. porriforius L. is cultivated in western Europe; its roots are 
used as a vegetable. Many of the species are used as food in the Caucasus 
and Soviet Central Asia. The roots can be used as a substitute for coffee. 
They contain a lot of inulin. 

Some Tragopogon species are melliferous, medicinal or fodder plants. 

Identification of the species is extremely difficult, as florets (for 
corolla color) and mature achenes are essential for precise identification. 

For the construction of the key and the preparation of the descriptions 
of the species, mainly the outer achenes and not the inner were 
considered. 

1. Florets yellow; peduncles below capitula strongly thickened; 
capitula large, 3.5-9 cm long 2. 

+ Florets yellow or pale yellow, dry florets sometimes lilac or reddish; 
peduncles weakly or strongly thickened or not thickened, but then 
florets violet or purple 8. 

2. Florets 3.5-6.5 cm long; achenes with pappus 3.5-5 cm, with thick, 
sulcate beak about 3 mm long, pappus longer than achene, 2.3- 
2.6 cm long, with hairy annulus at base (on sands of Soviet Central 

Asia and western Siberia); Section 12. Dubjanskya Boriss 

60. T. sabulosus Krasch. and S. Nikit. 

+ Achenes with slender, long beaks, slightly shorter, as long as, or 
exceeding body of achene; pappus shorter than body of achene or 
almost as long 3. 

3. Achenes weakly sulcate, smooth, gradually narrowed into long and 
slender beak as long as or slightly shorter than body of achene; 
involucral bracts 6-8 or 8-12(16) (European part, Soviet Central 
Asia); Section 1. Majores (Artemcz.) Kuthath 4. 

+ Achenes deeply sulcate, with acute, prominent, rugose ribs, abruptly 
narrowed into long and slender, curved, smooth beak, about 2 cm 
long, two times as long as body of achene; pappus about 2.5 cm 
long; involucral bracts eight, exceeding florets and achenes 

(Caucasus); Section 2. Kemularia Boriss 

6. T. kemulariae Kuthath. 

4. Achenes with pappus 6-8 cm long, smooth or very deeply rugose, 
obtusely five-angled, mostly whitish, with smooth slender beak 



118 

without clavate thickening at apex; pappus 3-3.5 cm long, almost 
as long as achene, excluding pappus; cauline leaves usually up to 
4-6 cm wide, sometimes much narrower, oblong-lanceolate to 
lanceolate; capitula 6.5-9 cm long; involucral bracts eight(nine); 

plants robust, 40-100 cm high (Pamiro-Alai) 

3. T. paradoxus S. Nikit. 

+ Achenes with pappus 4.8-5.5 cm long, weakly or more or less 
118 strongly rugose, sulcate and ribbed 5. 

5. Beak of achenes clavate, very thin, bent; achenes with acute ribs; 
pappus two-thirds as long as achene including beak; involucral 
bracts or equaling exceeding achene 7. 

+ Beak of achenes straight, not or weakly clavate; achenes 
weakly squamose-tuberculate and sulcate; pappus more or less 
as long as achene; involucral bracts six to eight, usually shorter 
than achenes 6. 

6. Achenes with pappus about 5.5 cm long; beak slender, 1.3-1.7 cm 
long; pappus about 2.5 cm long, whitish; involucral bracts 
(seven)eight, much shorter than achenes; capitula 6-7 cm long 
(Pamiro-Alai) 4. T. serawschanicus S. Nikit. 

+ Achenes with pappus about 4.8 cm long; beak somewhat thick, 
about 1.1 cm long, almost smooth; pappus about 2.3 cm long, 
yellowish; involucral six to eight, as long or slightly shorter 
than achenes, much longer than florets; capitula cylindrical, 3.5- 
5 cm long (Balkhash Region, Dzhungaria-Tarbagatai, Tien 
Shan) 5. T. pseudomajor S. Nikit. 

7. Achenes with pappus 3.2-5.5 cm long; beak 0.8-2 cm long; body 
of achenes weakly sulcate, tuberculate from acute imbricate scales, 
continued on lower part of beak; pappus dirty white or yellowish, 
1.2-2.5 cm long; involucral bracts (8)10-12(16), considerably 
longer than florets, shorter or more or less as long as achenes; 
capitula 5-6(7) cm long (European part) 1. T. dubius Scop. 

-I- Achenes with pappus 5.5-7.5 cm long; beak 1.7-2.5 cm long, thin, 
finely sulcate, smooth; body of achene indistinctly ribbed, weakly 
rugose from small scales or almost smooth; pappus yellowish- 
golden-yellow, lustrous, 2.5-3 cm long; involucral bracts 
considerably longer than florets and slightly longer or, less often, 
almost as long as achenes; capitula 5.5.-8.5 cm long at fruiting 

(Soviet Central Asia) 2. T. capitatus S. Nikit. 

8(1) Roots cylindrical, swollen, tuberous to some depth; florets pale- 
or bright-yellow, dry florets lilac, with brownish veins (Caucasus). 
Section 7. Tuberosi Boriss 9. 

-H Roots cylindrical but not tuberous; florets yellow, violet, pink, or 
purple 10. 



119 

9. Florets pale yellow; involucral bracts shorter than florets; capitula 
small and medium; peduncles not thickened; plants short, 5-15(20) 

cm high, floccose, mainly in lower part 

27. T. pusillus M.B. 

+ Rorets bright yellow, sometimes with violet veins; involucral bracts 

as long as or shorter than ligulate florets; capitula large, at fruiting 

119 4-5 cm long; peduncles thickened below capitula; plants 20-40(45) 

cm high 26. T. tuberosus C. Koch 

10. Pappus almost three times as long as achene with beak; beak long, 
one and one-half to two times as long as body of achene; involucral 
bracts eight; florets pale yellow (Caucasus); Section 4. Macropogon 
(Kuthath.) Boriss 10. T. macropogon C.A.M. 

+ Pappus shorter, as long as or slightly longer than body of achene; 
combination of characters different 11. 

1 1 . Peduncles strongly thickened below capitula, fistular; leaves linear- 
lanceolate, upward spreading; involucral bracts five to eight, as 
long as or longer than florets; achenes 5-8 cm long, rugose- 
tuberculate with thick straight beak, almost as long as or shorter 
than achenes. Mainly cultivated in western Europe, sometimes wild; 
Section 17. Hebecarpus Boriss 79. T. porrifolius L. 

+ Peduncles not thickened or weakly or strongly thickened, but then 
combination of characters different 12. 

12 Florets violet, purple or pink; beak somewhat thick, straight, as 
long as body of achene or shorter 13. 

+ Florets yellow, less often violet or purple, but then achenes with 
long slender beak 15. 

13 Pappus violet or reddish; involucral bracts six to eight; one and 
one-half to two times as long as florets; peduncles clavate, short; 
capitula large, 4-7 cm long (Caucasus); Section 15. Coloropappus 
Boriss 14. 

+ Pappus whitish or yellowish, less often faintly colored in upper 
part; involucral bracts 8-10, shorter or almost as long as florets; 
plants short, 5^0 cm high usually with leaves crowded in lower 
part of plant; peduncles not or slightly thickened, less often 
clavately thickened; capitula medium to large; plants mostly of 
Soviet Central Asia; one species in the Caucasus and one in the 

southeast of the European part; Section 16. Rubriflori Boiss 

69. 

14. Capitula 4-5.5 cm long at fruiting; achenes 4-5 cm long with 
pappus; pappus one and one-half to two times as long as achene, 
purple or reddish in upper part; body of achenes with five 
prominent squamose-tuberculate ribs, abruptly terminated into thick 
smooth beak, one and one-half times as long as achene; plants 



120 

40-60 cm high (Caucasus) 67. T. nachitschevanicus Kuthath. 

+ Capitula about 7 cm long at fruiting; achenes 4-5 cm long with 
pappus; pappus dirty violet, as long as or slightly longer than 
achenes; body of achene deeply sulcate, two times as long as 
120 somewhat thickened beak; plants 7-30(60) cm high (Caucasus).. 

66. T. coloratus C.A.M. 

15 Florets dark violet or reddish; peduncles weakly thickened below 
capitula; capitula 6-8 cm long at fruiting; achenes large, with long 
slender beak; pappus shorter than achene, whitish or yellowish; 
involucral bracts 7-8(12), almost two times as long as florets; large, 
sparingly branched, 30-120 cm high plants (Soviet Central Asia, 

Caucasus); Section 5. Krascheninnikovia Boriss 

11. T. krascheninnikovii S. Nikit. 

+ Florets yellow or reddish-violet, but combination of characters 
different 16. 

16. Achenes gradually narrowed into slender long beak as long as or 
slightly shorter than body of achene; florets yellow or reddish- 
violet 17. 

+ Achenes abruptly narrowed into beak, or achene without beak, or 
beak indistinct 20. 

17. Involucral bracts five to eight, exceeding florets; capitula narrow- 
cylindrical, with few seeds, about 5 cm long; perennial plants with 
narrowly-linear leaves (Soviet Central Asia, Caucasus); Section 3. 
Angustissimi Kuthath 18. 

+ Involucral bracts 6-11, usually 7-8, exceeding florets or shorter; 
capitula usually medium-sized, on long straight peduncles; perennial 
or biennial plants; Section 6. Tragopogon 36. 

18. Florets yellow; involucral bracts five to seven, two times as long 
as florets; achenes 3.5^ cm long 19. 

+ Florets reddish-violet or pink; involucral bracts eight, slightly 
exceeding florets; achenes with about 4.2 cm long pappus and 
about 6 mm long beak, clavately thickened below pappus; plants 
glaucescent-green 7. T. sosnovskyi Kuthath. 

19. Involucral bracts five; achenes with finely sulcate slender beak, 
4-7(9) mm long, clavately thickened at apex and sparsely pubescent 
but without hairy annulus at base of pappus; pappus about 2 cm 
long, sometimes brownish; plants glabrous, arachnoid-tomentose 
in lower part when young, strongly branched and densely leafy; 

leaves linear to filiform (Soviet Central Asia) 

9. T. vvedenskyi M. Pop. 

+ Involucral bracts five to seven; achenes with about 9 mm-long 
slender beak without thickening at apex and without hairy annulus; 
plant loosely floccose (Caucasus) 8. T. segetus Kuthatfi. 



121 

20(16) Achenes with five prominent winged ribs, abruptly narrowed into 
beak; pappus longer than achene; capitula large; involucral bracts 
8, less often 10-12, sometimes 5; biennial plants (Caucasus); 

Section 8. Profundisulcati Kuthath 26. 

+ Achenes without winged ribs, without or with indistinct beak; 
121 capitula medium to large, 2-5 cm long; involucral bracts 6-8 or 

8-12, sometimes five, but then achenes without beak 

21. 

21. Achenes somewhat thick, acutely tuberculate and sulcate, without 
beak; pappus longer than achene; involucral bracts five (less often 
six), shorter than florets and achenes; capitula numerous, small to 
medium, 2-3.5 cm long; florets pale yellow, dry florets pale lilac. 
Plants perennial or biennial, branched from base (Kopetdag Range); 

Section 13. Dasypogon Boriss 

61. T. kopetdaghensis Boriss. 

+ Achenes somewhat thick, smooth, without or with short thick beak, 
or achenes slender and smooth or rugose, gradually narrowed into 
indistinct beak; pappus shorter than or almost as long as achenes; 
capitula small to large; involucral bracts 6-12; florets yellow or 
pale yellow 22. 

22. Capitula 2^.5 cm long, numerous, on short usually bent peduncles, 
in paniculate inflorescence; achenes somewhat thick, without or 
with short thick beak, smooth; biennial plants (sands of Soviet 
Central Asia and lower reaches of the Volga and Don); Section 
14. Nikitinia Boriss 23. 

+ Capitula small and medium to large, on erect, mostly long 
peduncles; achenes smooth or shallow sulcate, gradually narrowed 
into indistinct, slender or somewhat thick beak; perennial and 
biennial plants 50. 

23. Leaves lanceolate, basally broad; plants large; capitula numerous, 
2.5-4.5 cm long; achenes thick, 2-3.5 cm long including pappus, 
smooth, finely sulcate; pappus about 1.5 cm long; plants 40-150 
cm high with strong up to 1.5 cm thick stem (sands of the Aral 
Region) 62. T. dubjanskyi Krasch. and S. Nikit. 

+ Leaves linear; plants flocculose-tomentose, sometimes only near 
leaf bases; capitula small to medium and large, 1-4 cm long; 
achenes rugose or with alternafing rugose and smooth edges, 
shallow-sulcate 24. 

24. Pappus without hairy annulus at base, about 1 cm Ipng; capitula 
1.5-2.5 cm long , 25. 

+ Pappus with hairy annulus at base, 1.5-1.8 cm long, longer than 
achenes; achenes somewhat thick, shallow-sulcate, with alternating 
smooth and rugose edges and hairy annulus at apex; capitula 20- 



122 

40 mm long; involucral bracts 7-12; plants glaucous from 

tomentum (Lower Volga, Lower Don, Aralo-Caspian Region) .... 

64. T. ruthenicus (C.A.M.) Bess. 

25. Plants up to 1.5 m high, almost glabrous and flocculose-tomentose 

only in upper part and near leaf base, strongly branched above; 

involucral bracts 8-10; cauline leaves about 8 mm wide at base, 

122 carinate 63. T. scoparius S. Nikit. 

+ Plants 10-40 cm high, more or less densely pubescent below 

capitula, young branches flocculose usually branched from base; 

involucral bracts (seven) eight, densely pubescent 

65. T. donetzicus Artemcz. 

26(20) Achenes gradually narrowed into beak; basal leaves mostly linear 

27. 

+ Achenes abruptly narrowed into beak; basal leaves lanceolate and 

broadly lanceolate 28. 

27. Basal and cauline leaves almost similar, linear or broadly linear, 
up to 1.5 cm wide; upper leaves reduced and broadened, at base 
up to 2 cm wide, lanceolate, all leaves dark green and lustrous; 

florets yellow, dry florets with light blue stripes 

37. T. makaschwilii Kuthath. 

+ Basal leaves narrowly linear, folded, long and subulate; florets 
yellow 35. T. acanthocarpus Boiss. 

28. Leaves 1.5^ cm wide 29. 

+ Leaves much narrower..... ...30. 

29. Involucral bracts 10-12, wide, lanceolate, long-acuminate, shorter 
than florets, basally flocculose at flowering; beak shorter than body 
of achene; pappus whitish 34. T. latifolius Boiss. 

+ Involucral bracts eight, narrowly lanceolate, white-membranous 
along margin; beak as long as body of achene, clavate; body of 
achene shallow-sulcate, white-scaly along ribs, with five obtuse 
edges; pappus yellowish-golden, shorter than achene including beak 
36. T. idae Kuthath. 

30. Involucral bracts 5 or 10-12 34. 

+ Involucral bracts eight 31. 

31. Peripheral achenes large, 4-6 cm long, with five deep furrows and 
five prominent winged ribs 32. 

+ Peripheral achenes smaller 33. 

32. Basal leaves long, almost half as long as plant, linear or broadly 
linear; plant glabrous; beak slender, more or less as long as achene, 
with hairy annulus at apex; pappus shorter than achene with beak; 

involucral bracts as long as or slightly shorter than achene 

29. T. ketzkhovelii Kuthath. 

+ Basal leaves 8-10 cm long, up to 3 cm wide, oblong-lanceolate; 



123 

beak somewhat thick, as long as or longer than body of achene 

30. T. plantagineus Boiss. and Huet 

33. Outer achenes with five deep furrows, acutely tuberculate- scaly 

123 along ribs, abruptly narrowed into beak longer than its body; 
involucral bracts as long as or longer than achenes; florets pale 

yellow (Southwestern Transcaucasia) 

28. T. meskheticus Kuthath. 

+ Outer achenes with 10 ribs, slender, sulcate, white scaly, bent, 
convex near midpoint, abruptly narrowed into beak as long as or 
shghtiy longer than its body; capitula large; involucral bracts eight, 
shorter than achene including pappus . 33. T. karjaginii Kuthath. 
34(30) Involucral bracts five; capitula large, about 4.5 cm long; marginal 
achenes with five deep furrows and distinct ribs; beak shorter than 
body of achene, thickened at apex; pappus almost as long as achene 
38. T. armeniacus Kuthath. 

+ Involucral bracts 10-12 35. 

35. Involucral bracts 10-12; basal leaves up to 23 cm long, 7-8 
mm wide. Cauline leaves shorter, with prominent veins; upper 

leaves short, up to 2-2.5 cm wide at base 

31. T. buphtalmoides Boiss. 

+ Involucral bracts 12; basal leaves linear, shorter, not exceeding 

stem; cauline leaves numerous, crowded, reduced, lanceolate 

32. T. tasch-kala Kuthath. 

36(17) Florets yellow, pale yellow or orange, sometimes dry florets pale 
lilac or whitish; pappus longer or shorter than achene 41. 

+ Florets violet, pink-violet, or pink; pappus shorter or as long as 
achenes with beak, sometimes even longer 37. 

37. Involucral bracts 8-12, exceeding florets; peduncles slightly 
thickened or not below capitula; biennial plants of Western Siberia 
and the Volga Region 40. 

+ Involucral bracts seven or eight, almost as long as florets; peduncles 
not thickened; perennial plants of Soviet Central Asia 38. 

38. Plants with downy white tomentum on stem, leaves, and involucral 
bracts; root collar with fibrous remnants of old leaves; pappus 
more or less 2.5 cm long, much longer than achene, brownish; 
beak somewhat slender, weakly rugose in lower part, pubescent 
above 21. T. tomentosulus Boriss. 

+ Plants glabrous; pappus shorter or almost as long as achene with 
beak 39. 

39. Plants 10-25 cm high, caespitose; florets pink- violet; achenes with 
pappus about 2.8 cm long; beak about 6 mm long, shorter than 

124 body of achene; pappus about 1.3 cm long (Tien Shan, Dzhungarian 
Alatau) 23. T. subalpinus S. Nikit. 



124 

+ Plants 25-30 cm high; florets pink; achenes with pappus about 4.8 
cm long; beak about 1.1 cm long, as long as body of achene; 

pappus about 2.3 cm long (Balkhash) 

20. T. karelinii S. Nikit. 

40. Florets dark red or purple, shorter than involucre; peduncles slightly 
thickened below capitula; plants 45-110 cm high; achenes with 
pappus about 4 cm long, smooth, terete, with slender smooth beak 
as long as, shorter, or slightly longer than body of achene; pappus 

about 1.7 cm long (Western Siberia) 

24. T. sibiricus Ganeschin 

+ Florets pink-violet or purple, aggregated in narrow capitula, 2.5- 
3.5(5.5) cm long; peduncles not or sUghtly thickened, glabrous, 
slender; leaves narrowly linear to filiform; achenes straight, five- 
angled, rugose in lower part, with slender beak longer than body 
of achene; pappus shorter than achene including beak, dirty brown 

(on chalk near Vol'sk on Volga) 25. T. cretaceus S. Nikit. 

41(36) In volucral bracts much shorter, two-thirds as long as outer florets; 
florets bright, golden-yellow; achenes including pappus about 3.9 
cm long; body of achene gradually narrowed into about 9 mm- 
long beak; pappus much shorter than achene with beak (European 
part) 12. T. orientalis L. 

+ Involucral bracts as long as florets or longer, less often slightly 
shorter. Florets pale yellow, yellow or orange; dry florets whitish 
or pale lilac 42. 

42. Florets orange; body of achene obtusely sulcate with alternating 
rugose and smooth edges, abruptly narrowed into 5-7 mm-long 
beak; involucral bracts divergent, as long as achenes and shorter 
than florets (Altai) 19. T. altaicus S. Nikit. and Schischk. 

+ Florets light yellow or yellow, sometimes dry florets whitish or 
pale lilac 43. 

43. Plants glabrous, green; involucral bracts almost as long as florets 
or longer 45. 

+ Plants whitish from flocculose tomentum, sometimes pubescent only 
in early developmental stage 44. 

44. Plants perennial, short, 10-20(25) cm high, caespitose, young 
branches flocculose-tomentose; involucral bracts six (less often 
seven or eight); florets yellow, dry florets pale lilac (Pamir, Alai 
Range) 22. T. gracilis D. Don. 

+ Plants biennial whitish from flocculose-tomentum; involucral bracts 
125 7-11; florets pale yellow, whitish on drying (Belorussia, sands).. 
14. T. bjelorussicus Artemcz. 

45. Florets light yellow above, dirty violet below; capitula 15-20 mm 
long (Dauria) 15. T. trachycarpus S. Nikit. 



125 

+ Florets yellow or light yellow; capitula 25-50 mm long (European 
part, Soviet Central Asia) 46. 

46. Plants perennial; root collar covered with fibers; peripheral achenes 

smooth; beak as long as body of achene (Lithuania) 

40. T. gorskianus Rchb. 

+ Plants biennial; root collar covered with fibers or old leaves . 47. 

47. Florets light yellow; root collar covered with fibers; involucral 

bracts 8-10; almost as long as florets (European part) 

13. T. pratensis L. 

+ Florets yellow; root collar with or without renmants of old leaves; 
involucral bracts seven or eight(nine), exceeding florets; montane 
plants 48. 

48. Peduncles at fruiting more or less thickened; achenes with pappus 
4-4.5 cm long, with 1-1.2 cm long beak, and hairy annulus at 
apex below pappus 49. 

+ Peduncles not thickened at fruiting; achenes with pappus 3.2-3.6 
cm long, with about 0.6 cm-long beak, and without hairy annulus 
at apex (north of Trans-Ili Alatau, Dzhungarian Alatau, Tarbagatai) 
16. T. songoricus S. Nikit. 

49. Plants (25)50-100 cm high, stem simple or branched, usually in 
upper part, sparsely leafy; involucral bracts seven or eight(nine), 
one and one-half times as long as florets; peduncles at fruiting 
slightly thickened; florets yellow; achenes about 4 cm long, with 
about 1 cm long beak (Tien Shan, Pamiro-Alai, at 1,000-2,600 m) 
17. T. turkestanicus S. Nikit. 

+ Plants 15-35 cm high, branched from base, densely leafy in lower 
part; involucral bracts eight(nine), slightly shorter or as long as 
florets; peduncles more or less thickened at fruiting; florets pale 
yellow, pale violet when dry; achenes about 4.3 cm long, with 
about 1.2 cm-long beak (at 3,200-4,000 m in the Gomo [Moun- 

tainous]-Badakhshan Autonomous Region) 

18. T. badachschanicus Boriss. 

50(22) Capitula small to medium, up to 3 cm long; florets yellow and 
pale yellow; involucral bracts six to eight or 8-12, shorter than 
florets and achenes; achenes small, slender, smooth or rugose 
and tuberculate, narrowed into short, indistinct, 1.5-3 mm-long 
beak 51. 

+ Capitula medium to large, 3.5-5 cm long; florets large, orange, 

yellow, or'pale yellow; involucral bracts eight, considerably shorter 

126 than florets and as long as achenes; achenes large, gradually 

narrowed into somewhat thickened, straight, indistinct beak shorter 

than the body (Caucasus); Section 11. Sosnovskya Boriss 52. 

5 1 . Achenes slender, tapered and smooth or weakly scabrous, narrowed 



126 

toward apex into indistinct, 1.5-3 mm-long beak; pappus shorter 
than achenes; involucral bracts 8-12; peduncles slightly longer than 
leaves; plants perennial, flocculose-tomentose, branched from base 
and often caespitose, less often plants biennial (sands, European 
part); Section 9. Bessera Boriss 55. 

+ Achenes shallow-sulcate, tuberculate, distinctly narrowed into short, 
sometimes indistinct, beak; pappus as long as achene; involucral 
bracts six to eight; peduncles longer than leaves; plants mostly 
perennial, less often biennial, not caespitose (European part, 

Caucasus, Turkmenia); Section 10. Brevirostres Kuthath 57. 

52(50) Achenes white, with indistinct, somewhat thickened beak shorter 
than the body; pappus longer than achene, about 2.5 cm, dirty 
golden; florets yellow; cauline leaves with white border (Caucasus) 
56. T. marginatus Boiss. and Huet 

+ Achenes gradually narrowed into 5-8(10) mm long beak; pappus 

almost as long as achene or shorter, white or grayish 53. 

53. Florets bright yellow, considerably longer than involucre; plants 
20-65 cm high; basal leaves linear; cauline leaves broadened from 
base, oblong, long-acuminate; usually all leaves flocculose, sinuate 
and drooping (Caucasus) 57. T. reticulatus Boiss. and Huet 

+ Florets pale yellow, sometimes dry florets pale lilac 54. 

54 Florets pale yellow; involucral bracts eight, lanceolate, shorter than 
achenes, at fruiting deflexed and involute; body of achene 
indistinctly ribbed in upper part, finely and indistinctly scaly along 

ribs; beak about 0.5 cm long (Caucasus) 

58. T. charadzeae Kuthath. 

+ Florets pale lilac; involucral bracts 8-10, much shorter than florets 
and achenes, lanceolate-linear, finely acuminate; body of achene 
sulcate, upper part squamose with acute, upward directed scales; 

beak 0.8-1.0 cm long (mountainous Turkmenia, Iran) 

59. T. maturatus Boriss. 

55(51) Plants perennial, 5-10 cm high, with strong shortly branched stem, 
caespitose, with numerous fertile and vegetative branches; leaves 
narrowly linear to filiform, numerous; capitula rather small; florets 
light yellow; achenes almost smooth; with about 3 mm-long, 

indistinct beak (Lithuania) 39. T. lithuanicus (DC). Boriss. 

127 + Plants biennial, 15-75 cm high, capitula 2.5-3.5 cm long; florets 
yellow or light yellow; achenes 2.5-3.5 cm long, slender, smooth 
or finely scarious-rugose along ribs, gradually and insignificantly 

narrowed toward apex; beak not developed 56. 

56. Florets yellow; involucral bracts (seven) eight, pubescent bracts 
alternating with less pubescent ones, shorter than florets; pappus 
9-10 mm long; ahcenes slender, five-angled, about 2.5 cm long, 



127 

finely scarious-rugose (sands along the middle reaches of the Don 
and Volga rivers) 41. T. tanaiticus Artemcz. 

+ Florets pale yellow; involucral bracts 8-12, with long cusp; pappus 
about 1.5 cm long, almost as long as achene; achenes 2.5-3.5 cm 
long, slender, almost smooth, in upper part narrowed over 2-3 

nmi (sands of the Baltic Region) 

42. T. heterospermus Schweigg. 

57(51) Plants perennial; beak of achenes much shorter tnan body, or 
achenes without beak; florets yellow or orange-yellow, dry florets 
sometimes with violet veins; capitula small to medium, up to 3 
cm long 58. 

+ Plants biennial; beak of achenes undeveloped or inconspicuous; 
florets yellow; capitula small 62. 

58. Florets orange-yellow, sometimes with purple stripes below ... 59. 
+ Florets yellow 60. 

59. Stems slender, numerous; plants glabrous; involucral bracts seven; 
achenes with pappus about 2 cm long; florets two tim«^a as long as 

involucre; orange-yellow (Western Caucasus) 

51. T. colchicus Alb. 

-I- Stems few, two or three; involucral bracts 8-10, flocculose at base; 
achenes with pappus about 2.4 cm long; florets slightly exceeding 
involucre, orange-yellow, with purple stripes below (South Osetia) 
50. T. filifolius Rehm. 

60. Root collar densely covered with fibers; beak slightly shorter or 
almost as long as body of achene, slender, distinct; florets pale 
yellow, whitish when dry; plants usually tall, 20-90 cm high, 

densely leafy; leaves 4-8 mm wide, linear to lanceolate 

47. T. graminifolius DC. 

-I- Root collar weakly covered with fibers or not; beak one-third as 
long as achenes; florets yellow, violet-lilac or pale yellow when 
dry; leaves narrowly linear, 1.5-4 mm wide (Caucasus) 61. 

61. Florets yellow, violet-lilac when dry; plants glabrous, grayish-green, 
sometimes arachnoid, flocculose up to flowering, 30-60(90) cm 
high; involucral bracts eight, lanceolate-linear; peripheral achenes 
2.5-2.8 cm long, whitish, shallow- sulcate, tuberculate-squamose, 

128 gradually narrowed into smooth, glabrous, 3-6 mm-long beak; 

pappus longer than achene 48. T. brevirostris DC. 

+ Florets pale yellow; plants sparsely flocculose, 35-50 cm high, 
involucral bracts six to eight, lanceolate; peripheral achenes about 
2.5 cm long, squamose-tuberculate, globose, and obtusely angular 
with two rows of sparse, indistinct scales; body of achene distinctly 
narrowed into about 0.5 mm-long beak; pappus shorter than achene 
49. T. serotinus Sosn. 



128 

62(57) Achenes with or without indistinct beak 63. 

+ Achenes with distinct short beak 66. 

63. Pappus 0.5-0.7 cm long, shorter than achene; achenes without or 
with indistinct beak 64. 

+ Pappus about 1.5 cm long, as long as achene or longer; achenes 
without beak 65. 

64. Achenes with 1.6-1.9 cm long pappus, without beak, smooth, 
slender; upper leaves strongly reduced, ovate-cordate at base (on 

sands in the lower reaches of the Dnieper and Bug rivers) 

55. T. borysthenicus Artemcz. 

+ Achenes with pappus 2-2.3 cm long, narrowed into indistinct, about 
3 mm-long beak; upper and cauline leaves acuminate (sands and 
sandy soils in the middle reaches of the Dnieper and Don rivers) 
53. T. ucrainicus Artemcz. 

65. Achenes with pappus about 2.3 cm long, curved, sulcate, distinctly 
narrowed toward apex; pappus about 1.3 cm; capitula rather small; 
plants arachnoid-tomentose, with narrow-linear leaves, usually 
convolute 52. T. stepposus (S. Nikit.) Stankov 

-I- Achenes with pappus about 3.2 cm long, straight, shallow-sulcate, 
with two rows of scales, elsewhere smooth, gradually narrowed 
toward apex; plants grayish from dense flocculose pubescence ... 

54. T. daghestanicus (Artemcz.) Kuthath. 

66(62) Basal leaves 5-6 cm wide (Crimea) 46. T. elatior Stev. 

-I- Basal leaves narrow, linear or linear-lanceolate 67. 

67. Pappus shorter than achene; basal leaves linear, not folded, usually 
flat; upper and middle cauline leaves broadened at base, finely 
acuminate, sometimes convolute 45. T. podolicus Bess. 

+ Pappus as long as or longer than achene 68. 

68. Plants glabrous or weakly pubescent, sometimes flocculose- 
tomentose in leaf axils; basal leaves linear, usually folded; 
involucral bracts (seven) eight, shorter than florets; capitula often 
solitary, on long, straight, somewhat slender peduncles; achenes 
with acute scales 43. T. dasyrrhynchus Artemcz. 

129 + Plants glabrous, light green; leaves not exceeding stem, herbaceous, 
linear or linear-lanceolate, flat; not broadened at base; involucral 
bracts five to eight, shorter than florets or as long, shorter than 
achenes; capitula narrow, small, 18-25(30) cm long; achenes finely 
tuberculate along ribs; sulcate 44. T. volgensis S. Nikit. 

69. Peduncles strongly clavately thickened below capitula, sulcate; 
leaves broadly lanceolate, 10-15(20) mm wide; involucral bracts 
eight, almost as long as florets; achenes about 3.6 cm long, with 

about 0.8 cm-long beak, without hairy annulus 

77. T. kultiassovii M. Pop. ex S. Nikit. 



129 

+ Peduncles below capitula not or weakly thickened 70. 

70. Pappus two-thirds as long as achene with beak, brownish, densely 
hairy; leaves with broad rhomboid base; stems slender, erect; 
achenes with pappus 3.5-3.7 cm long, deeply sulcate, sharply 
scabrous, abruptly narrowed into straight beak (Balkhash Region, 

Zaisan Kettle Hole, Dzhungarian Alatau) 

75. T. elongatus S. Nikit. 

+ Pappus as long as achene with beak or longer 71. 

7 1 . Plants tall, glabrous, 40-60 cm high, with narrowly linear to linear- 
lanceolate leaves; achenes 5-6 cm long, body of achene gradually 
narrowed into thick, straight, 0.7-1.2 cm-long beak, sulcate in lower 
part (Alai Range) 73. T. alaicus S. Nikit. 

-I- Plants shorter, 10^0 cm high, often with narrow or wide leaves 
crowded at base; plants glabrous or young branches flocculose; 
achenes 4-5 cm long 72. 

72. Plants glabrous, short, with short peduncles not thickened or 
somewhat thickened at fruiting 73. 

-I- Plants flocculose, especially near leaf bases, sometimes hairy only 
when young, later glabrous; peduncles not thickened; leaves 
crowded in lower part of stem 77. 

73. Leaves lanceolate or oblong-lanceolate, up to 1 cm wide; achenes 
with about 1 cm-long beak, as long as body of achene, with hairy 
annulus at apex below pappus; pappus longer than achene, 3 cm 

long; plants 10-25 cm high (Kopetdag) 

74. T. gaudanicus Boriss. 

+ Leaves narrowly linear or lanceolate 74. 

74. Beak of achenes about 5 mm long, without hairy annulus at apex; 
pappus 2.5 cm long, as long as achene with beak or longer; achene 
with distinct sculpture, obtusely short-scaly; leaves on short (10- 
17 cm high) plants narrowly-linear, folded and straight (Zeravshan 
Range) 71. T. conduplicatus S. Nikit. 

130 + Beak 1-1.5 cm long, somewhat thick, smooth, with hairy annulus 
at apex, or without annulus, but then leaves wider; achenes five- 
angled, sulcate, finely and acutely squamose 75. 

75. Leaves wide, flat, with many veins, distant, sometimes reduced, 
lower leaves may be linear; peduncles not or weakly thickened; 
capitula large, at fruiting 4.5-5.5(6) cm long; involucral bracts 
shorter or almost as long as florets; beak about 1 cm long; pappus 

as long as achene with beak 

72. T. montanus S. Nikit. 

-I- Leaves linear, not longer than inflorescence; peduncles more or 
less thickened below capitula 76. 

76. Pappus brownish, densely plumose-villous, slightly shorter than 



130 

achene; leaves not longer than inflorescence, linear, sinuate; plants 
densely leafy (Karatau) 76. T. malicus S. Nikit. 

+ Pappus dirty white or yellowish, almost as long as achene; leaves 
broadened near base up to 5-12(20) mm, semiamplexicaul, 
narrowly linear or linear-lanceolate upward, narrowly membranous 

along margin (sandy deserts of central Kazakhstan) 

78. T. kasachstanicus S. Nikit. 

77(72) Beak of peripheral achenes gradually merging with body of achene 
78. 

+ Beak of peripheral achenes more or less abruptly joined with 
broader, deeply sulcate, sometimes winged part of achene; leaves 
10-20(30) mm wide, with cartilaginous border, undulate; plants 

10-40 cm high, densely leafy 70. T. marginifolius Pavi. 

78. Beak of achenes smooth; achenes weakly scabrous in upper part, 
more or less smooth below; basal leaves numerous, lanceolate- 
linear to filiform, flat or convolute (Caucasus) 

68. T. collinus DC. 

+ Beak, as also entire achene, sulcate, scabrous; leaves usually linear 
(Lower Volga, Aralo-Caspian) 69. T. ruber S.G. Gmel. 

Section 1. Majores (Artemcz.) Kuthath. Kavk. Predst. Tragop. (1957) 
37 emend. — Ser. Majores Artemcz. in Uch. Zap. Chernov. Univ. I (1948) 
88, diagnosis in Russian. — Peduncles below capitula mostly strongly 
thickened, less often weakly thickened; capitula large or medium, solitary, 
on long straight peduncles; involucral bracts 8-12 or 6-8, longer than 
ligulate florets; florets yellow, sometimes whitish or pale lilac on drying; 
achenes large, body of achene shallow-sulcate, finely rugose or smooth, 
gradually narrowed into long, slender, smooth beak, often longer than 
131 body of achene; pappus usually shorter or almost as long as achene 
with beak; plants biennial. 

Type of section: T. dubius Scop. 

Series 1. Robusti Kuthath. Kavk. Predst. Tragop. (1957) 38, p. p.— 
Peduncles thickened below capitula, fistular; capitula large; involucral 
bracts longer than ligulate florets; florets yellow; beak of achenes long, 
slender, longer than body of achene. 

1. T. dubius Scop. Fl. camiol. II (1772) 95; Montgomeri in Rhodora 
55 (1953) 659; Kuthath. Kavk. Predst. Tragop. (1957) 38; Fl. Azerb. 
VIII (1961) 507.— r. majus Jacq. Fl. Austr. I (1773) 19, t. 29; M.B. R 
taur.-cauc. II, 233, and III, 521; DC. Prodr. VII, 112; Ldb. Fl. Ross. II, 
784; Boiss. F. or. II, 747; Kryl. Fl. Zap. Sib. XI, 2975; Artemcz. in Tr. 
Inst. Bot. Khar'k. Univ. II, 55; Grossh. Fl. Kavk. IV (1934) 230, p. p.; 



131 

Klokov in Vizn. Rosl. URSR, 591; Kuthath. in Fl. Gruzii VIII, 620; 
Mikhail, in Fl. BSSR, V, 178; C. Rgl. in Berichte Schweiz. Bot. 
Gesellsch. 65 (1955) Karte 2. — T. desertorum Lindem. Fl. Cherson. I 
(1881) 365; Klokov, op. cit. 591. — T. major Jacq. var. vulgare Schmalh. 
Fl. II (1896) 146.— r. livescens Bess. Enum. pi. Volh. (1822) 30; DC. 
Prodr. VII, 112; Klokov, op. cit. 590. — T. intermedius Bess. op. cit. 83; 
DC. op. cit. 114, pro syn. — T. campestris Bess. Enum. pi. Volh. (1822) 
84; DC. op. cit. 114; Ldb. Fl. Ross. II, 784.-7. major Jacq. var. 
campestris (Bess.) Schmalh. op. cit.; Grossh. op. cit. — T. majus Jacq. (3. 
stenophyllus Boiss. I.e. — T. dubius Scop. II. campestris (Bess.) Hayek. 
Prodr. Fl. Balk. 2 (1931) 819.— Ic: Javorka and Csapody, Iconogr. Fl. 
Hung. (1929) f. 3991; Hegi, 111. Fl. VI, 2, fig. 733; Kuthath. op. cit. 
Plate VI, Fig. 21, Map No. 7. 

Biennial. Root vertical, easily pulled; root collar covered with 
previous year's leaves or not; plants 30-70 cm, sometimes 1.5 m high, 
with erect, simple or weakly branched, finely ribbed stem, glabrous or 
sometimes with weak flocculose pubescence at leaf base and below 
capitula. Lower leaves narrowly linear or lanceolate-linear, numerous, 
flat, erect, sometimes withering early; middle leaves (6)8-20 cm long, 
5-20 mm wide, broadened at base, semiamplexicaul, abruptly narrowed 
into linear or narrowly lanceolate, sometimes folded lamina; upper leaves 
shorter, long-attenuate-acuminate, often reduced. Peduncles long, fistular, 
clavately thickened below capitula, especially at fruiting, 6-10(12) nmi 
thick, sometimes tomentose. Capitula terminal on stem and branches, 
large, at fruiting 5-6(7) cm long, 2-2.5 cm wide; involucre of 8-12 
(less often 16) bracts; involucral bracts linear-lanceolate, long- and finely 
acuminate, glabrous, much longer than florets, shorter or almost as long 
as achenes, 4.5-6(7) cm long, 1.5-2 cm wide; florets yellow; achenes 
with pappus 3.2-5.5 cm long, without pappus 2-3 cm long, without 
beak 1-1.2 cm long, 1-1.2 mm wide, oblong, slightly curved, grayish, 
132 yellowish, or light brown, shallow-sulcate, five-angled, densely squamose, 
serrate-dentate, gradually narrowed into slender, smooth or weakly sulcate 
beak below, 0.8-2 cm, almost as long as body of achene, or longer, 
slightly broadened below pappus and with hairy annulus; pappus whitish, 
dirty white or yellowish, 1 .2-2.5 cm long, with hairs of variable length. 
2n = 12. Flowering May to August; fruiting June to September. 

Dry meadows, scrubs, forest edges, as a weed of roadsides, cultivated 
fields, sandy and clayey desert-steppes, sometimes on sands. — European 
Part: Ladoga-Ilmen, Upper Volga, Volga-Kama, Upper Dnieper, Middle 
Dnieper, Volga-Don, Trans- Volga, Upper Dniester, Bessarabia, Lower 
Don, Black Sea Region, Crimea, Lower Volga; Caucasus:. Ciscaucasia, 
Dagestan, Eastern and Southern Transcaucasia, Talysh. General 
distribution: Central Europe, Atlantic Europe, Mediterranean, Balkans- 



132 

Asia Minor, Armenia and Kurdistan. Described from western Europe. 
Type in Italy. 

Note. T. dubius Scop, is a polymorphic, widely distributed species. 
Besides the typical form, var. campestre (Bess.) Schmalh., with an 
involucre of eight bracts, less thickened peduncles and narrower and 
smaller capitula, is also found; the narrow-leaved form, var. stenophyllus 
Boiss., has been reported from Crimea. 

2. T. capitatus S. Nikit. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk 
SSSR, VII, 12 (1937) 257; Boris, in Fl. Turkm. VII, 293; Kovalevskaja 
in Fl. Uzb. VI, 434. — T. majus auct. fl. turkest. ex parte, non Jacq. — T. 
macrocephalum S. Nikit. in Zakir. Fl. i. Rast. Zeravsh. 2 (1962) 419, 
nomen. 

Biennial. Root vertical, robust; root collar covered with numerous 
basal leaves. Plant large (30-100 cm high), glabrous, usually branched 
below middle. Stem thick, robust, sulcate. Cauline leaves linear- 
lanceolate, basally broadened to 10-20 mm, semiaplexicaul, abruptly 
narrowed upward; lower leaves linear. Peduncles below capitula clavate 
to 7-15 mm, fistular, long, exceeding inflorescence. Capitula large, with 
fruits 5.5-8.5 cm long, 1.7-3 cm wide. Involucre of 10-12 (less often 
8-14) bracts, far exceeding florets and slightly longer or very rarely 
almost as long as achenes; involucral bracts linear-lanceolate, long- and 
finely acuminate. Florets yellow; outer achenes 5.5-7.5 cm long, without 
pappus 2.8-3.8 cm long, about 1.5 mm wide, shghtly curved, indistinctly 
five-angled, almost smooth, with inconspicuous scales along edges, 
obliquely truncate near base, finely sulcate, gradually narrowed into 
filiform beak about 1.7-2.5 cm long, thickened above and with tuft of 
hairs; pappus 2.5-3 cm long, yellowish golden, lustrous, with prominent, 
134 sparse, longer hairs, shorter than achenes with beak; inner achenes weakly 
rugose and almost smooth. Flowering May to June; fruiting June to 
August. (Plate VII, Fig. 2.) 

Dry slopes in the foothills and middle mountain zone, bottomlands, 
scrubs, steppes and as weed in crops. — Soviet Central Asia: Balkhash 
Region, Aralo-Caspian, Syr-Darya, Pamiro-Alai, Tien Shan, mountainous 
Turkmenia. General distribution: Possibly distributed in neighboring parts 
of Iran and Afghanistan. Described from Katta-Karamuk River in 
Kirgizia. Type in Leningrad. 

Note. Differs from T. dubius Scop, by having larger achenes with a 
very long and thin beak, peduncles at fruiting very strongly clavate, up 
to 1.5 cm wide, and larger capitula. 

Series 2. Paradoxi Boriss. Leaves linear-lanceolate to broadly oblong, 
up to 4 cm wide; peduncles clavate; capitula large; florets yellow, whitish 
or pale lilac on drying; achenes with weakly sculptured, almost smooth. 



133 




133 Plate VH. 

1 — Tragopogon heterospermus Schweigg.; 2 — T. capitatus S. Nikit.; 
3 — T. sibiricus Ganesch. 



134 

five-angled, with long, slender, and smooth beak, not or indistinctly 
thickened at apex, with annulus of hairs near pappus base. 

3. T. paradoxus S. Nikit. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 
1, I (1933) 200; Boriss. in Fl. Turkm. VII, 292; Kovalevskaja in Fl. 
Uzb. VI, 434. — T. maracandicus M. Pop. in Tr. Uzb. Gos. Univ. N. S. 
27; Biol. 14 (1941) 95.— Ic: Fl. Turkm. VII, Plate XLIV, Fig. 1. 

Biennial or perennial. Root cordlike, thick, vertical; root collar 
covered with remnants of old leaves. Plant large, glabrous, 40-100 cm 
high, more or less branched from base. Stem usually robust, densely 
leafy, especially at base. Leaves linear-lanceolate to oblong-lanceolate 
and oblong, usually up to 4 cm wide, semiamplexicaul, acuminate. 
Peduncles clavate below capitula at fruiting, 8-15 mm thick. Capitula 
large, 6.5-9 cm long at fruiting, with 50-60 achenes. Involucral bracts 
eight or nine, lanceolate-linear or linear, acuminate, up to 10 mm wide 
at base, slightly longer than florets, shorter or almost as long as achenes, 
sometimes membranous along margin in lower part. Florets yellow, 
whitish or pale lilac on drying. Peripheral achenes with pappus 7-7.5(8) 
cm long; without pappus 3.5-4(5.5) cm long, whitish; body of achene 
1.6 cm long, about 2 mm wide, slightly concave, five-angled, somewhat 
attenuate at base and obliquely truncate, smooth, sometimes weakly 
scabrous-tuberculate along ribs in upper part, abruptly narrowed into 
slender, 2 cm long, finely sulcate, smooth beak, at apex only slightly 
and briefly broadened, with tuft of dense fluff; beak longer or almost as 
long as body of achene; pappus about 3.5 cm long, with unequal hairs, 
dirty white or light colored, somewhat shorter than achene with beak; 
inner achenes edged, weakly ribbed, smooth or very weakly sculptured 
in upper part. Flowering April to June: fruiting May to July. 
135 In mountains near the lower boundary of the juniper forest zone, at 

zone of contact with herb-feathergrass steppe, on stony and clayey slopes 
at 800-2,500 m. — Soviet Central Asia: Pamiro-Alai, mountainous 
Turkmenia (Kugitang Range). General distribution: Possibly distributed 
in neighboring areas of Iran and Afghanistan. Described from vicinity 
of Baldzhuan. Type in Leningrad. 

Note. T. paradoxus S. Nikit. is well distinguished from other species 
by the weak sculpturing of the large achenes. Very variable in leaf 
width and length. 

4. T. serawschanicus S. Nikit. in Tr. Bot. Inst. Akad. Nauk SSSR, 
1, I (1933) 200; Kovalevskaja in Fl. Uzb. VI, 433. 

Perennial or biennial. Root woody, robust, vertical; root collar more 
or less without leaves or with isolated old leaves. Plant glabrous, 25-50 
cm high. Stems somewhat slender, usually simple or branched almost 



135 

from base, usually few (two or three) from a single root. Cauline leaves 
broadly lanceolate, about 10 cm long, 1-2 cm wide, slightly naiTOwed 
toward base, semiamplexicaul, acute, but not acuminate; upper leaves 
reduced, peduncles 10-20 cm long, exceeding leaves, slightly thickened 
at fruiting, 2-5 mm thick. Capitula 6-7 cm long, few, usually one or 
two, with 11-25 achenes. Involucral bracts (seven) eight, linear- 
lanceolate, long-acuminate, shorter than achenes by 1-2 cm; florets 
yellow. Peripheral achenes with pappus about 5.5 cm long, without 
pappus 3-3.6(4) cm; achenes without beak about 1.7-2.3 cm long, 2 
mm wide, five-angled, ribbed, finely squamose-tuberculate, usually 
slightly attenuate in lower part, distinctly narrowed into long and slender 
1.3-1.7 cm long beak sUghtly shorter than thicker part [body] of achene, 
smooth, not thickened at apex, with a dense annulus of hairs; pappus 
whitish, 2.5 cm long, shorter than achene with beak. Flowering July; 
fruiting August. 

Mountain slopes. — Soviet Central Asia: Pamiro-Alai. Endemic. 
Described from Zeravshan Range. Type in Leningrad. 

5. T. pseudomajor S. Nikit. in Bot. Mat. Gerb. Bot. Inst. Akad. 
Nauk SSSR VII, 12 (1937) 258.-7. majus auct. fl. turk. et fl. ross. or. 
ex parte, non Jacq. 

Biennial. Root vertical, somewhat slender or robust; root collar with 
remnants of old dry leaves. Plant glabrous, 25-80(100) cm high. Stem 
branched from middle or simple, rather densely leafy. Lower leaves 
linear-lanceolate or linear, basally broad, semiamplexicaul; upper leaves 
oblong-ovate from base or oblong-lanceolate, 10-20 mm wide at base, 
gradually narrowed into narrow, acuminate, sometimes filiform lamina. 
Peduncles of fertile capitula clavate, 5-10 mm thick, fistular, usually 
short, almost as long as inflorescence or much longer. Capitula at fruiting 
cylindrical, 3.5-5 cm long, 1-3 cm wide. Involucral bracts six to eight, 
136 as long as or shorter than achenes and far exceeding florets; florets 
yellow. Peripheral achenes with pappus about 4.8 cm long, without 
pappus 2.5 cm long; achenes excluding beak about 1.4 cm long, about 
2 nrni wide, straight, distinctly five-angled in cross section, finely and 
acutely squamose, weakly sulcate, rather distinctly narrowed into straight, 
somewhat thick or slender beak which is shorter than body of achene, 
about 1.1 cm long, more or less glabrous, with hairy annulus at apex 
and practically not thickened or slightly thickened; pappus about 2.3 
cm long, almost as long as achene or slightly shorter, yellowish. 
Flowering June to July; fruiting July to August. 

Grassy, meadow-covered slopes of mountains, in river valleys and 
fields at 1,200-2,200 m. — Soviet Central Asia: Aralo-Caspian (eastern 
part), Balkhash Region, Dzhungaria-Tarbagatai, Syr-Darya, Tien Shan, 



136 

Pamiro-Alai. General distribution: Dzhungaria-Kashgaria. Described from 
Talass Alatau. Type in Leningrad. 

Note. It differs from T. dubius Scop, by the narrower cylindrical 
capitula with six to eight involucral bracts (and not 10-12), achenes 
shortly and abruptly narrowed at the apex, thick stems, peduncles strongly 
thickened below the capitula and quite short, large capitula and by the 
form of the leaves. From T. capitatus S. Nikit. it is distinguished by the 
size and structure of the achene, etuberculate and quite thick, straight 
beak (and not with a clearly developed white tubercle at the apex with 
a long, slender beak, which exceeds the length of the achene), as well 
as involucral bracts that are shorter than the achenes. A fodder plant. 

Section 2. Kemularia Boriss. sect, nova in Addenda XXVIII, 723. 
— Peduncles strongly thickened; involucral bracts eight; capitula large; 
florets yellow, shorter than involucral bracts; beak of achenes about 2 
cm long, slender, bent, smooth, clavate at apex, abruptly joined with 
deeply sulcate body of achene with acute prominent rugose ribs; pappus 
shorter than achene with beak; plant biennial. 

Type of section: T. kemulariae Kuthath. 

6. T. kemulariae Kuthath. in Zam. po Sist. i. Geogr. Rast. Tbil. 17 
(1953) 28; Kuthath. Fl. Gruzii VIII, 623; Kavk. Predst. Tragop. (1957) 
60; Fl. Azerb. VIII, 512.— Ic: Kuthath. (1953) Fig. 1; op. cit. (1957) 
Plate IX, Fig. 31, Map No. 10. 

Biennial. Root cordlike, cylindrical, straight; root collar without old 
leaves. Plants more or less flocculose and weakly pubescent; stem erect, 
cylindrical, finely sulcate, 35-60 cm high, simple or sparingly branched, 
with single capitulum each. Basal leaves linear, often folded, sometimes 
with fine white border along margin, long, but not exceeding stems; 
cauline leaves to linear, broadened from base, gradually acuminate toward 
apex; upper leaves semiamplexicaul, not exceeding inflorescence, finely 
acuminate, broadened from base. Peduncles strongly thickened below 
capitula. Capitula up to 6-8 cm long; florets yellow, shorter than 
137 involucre; involucral bracts eight, linear, gradually narrowed from broad 
base to filiform apex, exceeding florets and achenes, 6-8 cm long. 
Peripheral achenes with pappus about 5 cm long, without pappus 3.1 
cm long, curved; body of achene 1 . 1 cm long, five-angled, with sharply 
projecting ribs and five rather deep furrows, ribs with indistinct sculpture 
in the form of somewhat acute scales, abruptly narrowed into filiform, 
slender, bent and smooth, about 2 cm-long beak, clavate at apex and 
with hairy annulus; pappus about 2.5 cm long, golden-yellow; inner 
achenes up to 5.8 cm long and with up to 2.4 cm-long beak, with 
smooth or weakly rugose body of achene along ribs, basally almost 



137 

smooth; pappus shorter than achene with beak. Flowering May to July; 
fruiting July to August. 

Subalpine zone, in dry areas at 1,700-1,990 m. — Caucasus: Eastern 
Trancaucasia (Georgia, Dzhavakhetia, Azerbaidzhan), Southern 
Transcaucasia, Talysh. Described from region between villages of 
Bezhano and Khumrisi. Type in Tbilisi. 

Section 3. Angustissimi Kuthath. Kavk. Predst. Tragop. (1957) 34. 
Peduncles gradually and slightly thickened below capitula, straight; 
capitula narrowly cylindrical, large, about 5 cm long; involucral bracts 
five to seven (eight), longer than ligulate florets; florets yellow or reddish- 
violet. Achenes gradually narrowed into slender beak, shorter than body 
of achene; pappus almost as long as achene (slightly longer or somewhat 
shorter); plant biennial. 

Type of section: T. sosnovskyi Kuthath. 

7. T. sosnovskyi Kuthath. in Zam. po Sist. i Geogr. Rast. Tbil. 15 
(1949) 94; Kuthath. Kavk. Predst. Tragop. (1957) 35; Kuthath. in Fl. 
Azerb. Vlll, 506. — T. crocifolium auct. non L.: C.A. Mey. Verzeichn. 
(1931) 61; Ldb. Fl. Ross. II (i844-1846) 788, quoad pi. Talusch.— 7. 
angustissimus auct. non S. Nikit.; Grossh. Fl. Kavk. IV (1934) 231. — T. 
vvedenskyi auct. non M. Pop. ex PavL; Grossh. Opred. Rast. Kavk. (1949) 
507.— Ic: Kuthath. op. cit. (1957) Plate IV, Fig. 14, Map No. 6. 

Biennial. Root straight or sinuate, slightly thickened; root collar 
usually with remnants of previous year's leaves. Plants aeruginous, weakly 
flocculose at stem base. Stem 30-50 cm high, erect, solitary, angular, 
branched and thickened in forks. Leaves narrowly linear to filiform, 
long; cauline leaves shorter, broadened at base, semiamplexicaul, 
uppermost leaves very short, abruptly narrowed and acuminate. Capitula 
solitary, terminal on branches, rather large, about 5 cm long, cylindrical; 
peduncles gradually thickened below capitula, especially at fruiting. 
Involucral bracts five to eight, narrowly lanceolate, exceeding florets, 
as long as achenes with pappus or longer, weakly pubescent at base or 
138 glabrous; florets reddish violet. Achenes with pappus about 4.2 cm and 
without pappus 2 cm long, 1.5 mm wide, without beak 1.4 cm long, 
finely tuberculate-squamose and sulcate, gradually narrowed into slender, 
about 0.6 cm long, glabrous and clavate beak at apex; pappus dirty 
white, about 2.2 cm long, without hairy annulus at base. Flowering 
April to May; fruiting June to July. 

Dry stony and cobbly slopes, in mountain-steppe communities, to 
upper mountain zone. — Caucasus: Eastern and Southern Transcaucasia, 
Talysh. Endemic. Described from Armenia. Type in Tbilisi. 

Note. Based on the achenes, it is similar to T. vvedenskyi M. Pop. 



138 

ex Pavl. But differs from the latter by its red-violet florets (and not 
yellow), shorter involucral bracts, and peduncles thickened below the 
capitula and by the range and ecology. 

8. T. segetus Kuthath. in Zam. po Sist. i. Geogr. Rast. Tbil. 17 
(1953) 33; Kuthath. Kavk. Predst. Tragop. (1957) 37.— Ic: Kuthath. 
ibid. (1957) Plate IV, Fig. 15 (incorrect). Map No. 6. 

Biennial. Root thick, vertical; root collar with remnants of leaves. 
Plants loosely flocculose, up to 45 cm high. Stem erect, solitary, very 
densely leafy, strongly branched above, nodose at base of branches, 
flocculose at nodes. Basal leaves narrowly linear to filiform, folded, 
long; cauline leaves exceeding capitula. Peduncles thickened below 
capitula, weakly pubescent; capitula about 4 cm long. Involucral bracts 
five to seven, longer than florets, as long as achene with pappus at 
fruiting, narrowly lanceolate, long-acuminate, broadened toward base; 
florets yellow. Achenes with pappus 3.5-4 cm long, without pappus 
about 2 cm long and 1 mm wide, more or less distinctly ribbed, whitish, 
shallow-sulcate, glabrous; beak slender, about 0.9 cm long, without 
thickening and hairy annulus at apex; pappus about 1.8 cm long, shorter 
or as long as achenes. Flowering June. 

Wheat fields in the lower and middle mountain zones. — Caucasus: 
Southern Transcaucasia. Endemic. Described from Armenia. Type in 
Tbilisi; isotype in Leningrad. 

Note. It differs from the closely related species T. sosnovskyi 
Kuthath. by having a densely leafy stem, larger capitula, thicker 
peduncles, yellow florets, and deeply sulcate, ribbed, white achenes. 

9. T. vvedenskyi M. Pop. ex Pavl. in Byull. Mosk. Obshch. Isp. 
Prir. XLII (11 XII 1933) 138; Boriss. in Fl. Turkm. VII (1960) 294; 
Kovalevskaja in Fl. Uzb. VI, 432. — T. angustissimus S. Nikit. in Tr. 
Bot. Inst. Akad. Nauk SSSR, Ser. 1, I (28 XII 1933) 194.— Ic: Pavlov, 
op. cit. Fig. 4. 

Biennial. Root cordlike, rather short, 5-7 mm thick; root collar with 
remnants of previous year's leaves. Plant smooth, glaucous, usually 
arachnoid-tomentose below when young, erect, with slender stem, usually 
strongly branched and densely leafy, 30-60 cm high. Lower leaves 
139 numerous, very thin, glabrous; cauline leaves herbaceous, narrowed from 
weakly broadened base into narrowly linear, sometimes filiform, 2-5 
mm-wide lamina folded along keel; upper leaves usually reduced, as 
also middle cauline leaves, not exceeding inflorescence. Peduncles in 
fertile plants 1-6 cm long, 2-4 mm thick, slightly thickened below 
capitula; fertile capitula narrowly cylindrical, 40-50 nmi long, (5)10 
mm wide. Involucre almost two times as long as florets, slightly 



139 

exceeding achenes with pappus or as long; involucral bracts narrowly 
lanceolate, five; florets yellow. Achenes 10-18 (up to 30) in a capitulum; 
peripheral achenes more or less straight or slightly bent, with pappus 
about 3.6 cm long, without pappus 1.8-2.1(2.4) cm long, 1.5 mm wide 
in body; achenes excluding beak about 1.1 cm long, indistinctly obtusely 
five-angled, finely sulcate, finely white-tuberculate or squamose along 
ribs, gradually narrowed into finely sulcate, white, 4-7(9) mm long beak, 
inflated at apex and sparsely hairy, but without hairy annulus at pappus 
base; pappus brownish-white, sometimes reddish, about 2 cm long, almost 
as long as achene with beak or slightly longer; inner achenes smooth in 
lower part, with distant scales above, indistinctly sculptured. Flowering 
May to June; fruiting May to August. 

Plants widely distributed on mountain-steppe grassy and stony slopes, 
at bottom of ravines, from forest zone to subalpine zone, at 1,400- 
2,000 m. — Soviet Central Asia: Tien Shan (Talass Alatau, very rarely 
in southwestern part of Karatau Range), Pamiro-Alai (Zeravshan, Hissar, 
Turkestan and Alai ranges, Kugitang), mountainous Turkmenia 
(Kopetdag). Endemic. Described from Boroldai Mountain in Karatau 
Range. Type in Moscow; isotype in Leningrad. 

Note. In the Kopetdag, var. turcomanicus Boriss. occurs, which is 
distinguished by a longer beak (up to 9 mm long) and slightly larger 
capitula. 

Section 4. Macropogon (Kuthath.) Boriss. comb. nov. emend. — 
Ser. Macropogon Kuthath. Kavk. Predst. Tragop. (1957) 49, p. p. — 
Peduncles slightly thickened below capitula at fruiting; capitula solitary, 
4-5 cm long at fruiting, on straight long peduncles; involucral bracts 
eight, shorter than achenes and slightly longer than florets; florets pale 
yellow; body of achene deeply sulcate and densely tuberculate, 6-7 mm 
long, abruptly narrowed into slender, long beak which is one and one- 
half times as long as body; pappus almost three times as long as achene 
with beak; plants biennial. 

Type of section: T. macropogon C.A. Mey. 

10. T. macropogon C.A. Mey. in Bull. Soc. Nat. Mosc. XI (1838) 
273; Ldb. Fl. Ross. II, 785; Kuthath. Kavk. Predst. Tragop. (1957) 49; 
Kuthath. Fl. Azerb. VIII, 510.-7. major auct. non Jacq.: C.A. Mey. 
Verzeichn. (1831) 61, excl. syn.; Kuthath. in Fl. Gruzii VIII, 620, p. 
140 p. — T. majus auct. non. Jacq.: Boiss. Fl. or. Ill (1875) 747, non Jacq. p. 
p. — Ic: Kuthath. op. cit. Plate VII, Fig. 22, Map 9. 

Biennial. Root vertical, almost fusiform, brown, rather slender; root 
collar smooth or with remnants of dry leaves. Plant glabrous, 25-50 cm 
high; stem simple, strawlike, cylindrical, glabrous or almost glabrous. 



140 

angular, branched almost from base, with long branches, densely leafy. 
Leaves upright, straight, linear-lanceolate, glaucescent, glabrous or almost 
glabrous, not bent at apex, gradually acuminate, not broadened at base 
and not semiamplexicaul, 10-12 cm long, 4-6 mm wide, flat; basal and 
cauline leaves not exceeding inflorescence. Peduncles fistular, somewhat 
thickened at fruiting below capitula. Involucral bracts eight, glabrous, 
lanceolate-linear, long-acuminate, at flowering 3-3.5 cm long, somewhat 
longer than florets, about 5 cm long at fruiting, 3-4 nmi wide at base, 
flat, shorter than achenes; florets pale yellow, shorter than involucral 
bracts, outer florets 2.5-3 cm long. Achenes with pappus 4.5-5 cm long, 
without pappus 1.3-2 cm long; body of achene 0.6-0.7 cm long, about 
1 mm wide, sulcate, densely squamose-tuberculate, with 10 ribs, abruptly 
narrowed into slender 0.7-1.2 cm-long beak, finely sulcate, apically with 
hairy annulus below pappus and usually not inflated; pappus 3.5-3.8 
cm long, almost three times as long as achene with beak, golden-yellow, 
lustrous, with plumose simple hairs at apex; inner achenes longer, with 
longer beaks. Flowering May to June; fruiting June (Plate VIII, Fig. 1). 
Meadows and sandy places along seacoasts. — Caucasus: Eastern 
Trnscaucasia, Talysh. Endemic. Described from Talysh. Type in 
Leningrad. 

Section 5. Krascheninnikovia Boriss. sect. nov. in Addenda XXVIII, 
723.— Ser. Robusti Kuthath. Kavk. Predst. Tragop. (1957) 38, p. p.— 
Peduncles slightly thickened below capitula, straight; capitula large, 6- 
8 cm long, cylindrical; involucral bracts seven or eight, less often 12, 
far exceeding florets; florets dark violet or raspberry-colored; achenes 
large, with long and slender bent beak, gradually merging with finely 
sulcate and acutely tuberculate body of achene; plants biennial. 

Type of section: T. krascheninnikovii S. Nikit. 

Series 1. Longirostres Boriss. — Beak slender, bent, longer than body 
of achene. 

11. T. krascheninnikovii S. Nikit. in Tr. Bot. Inst. Akad. Nauk 
SSSR, Ser. 1, I (1933) 191; Boriss. in Fl. Turkm. VII, 297; Kuthath. in 
Fl. Azerb. VIII, 508; Kovalevskaja in Fl. Uzb. VI, 431.-— r. longirostris 
auct. fl. turk. et cauc. non Bischof. — T. clavulatum S. Nikit. in Bot. 
Mat. Gerb. Bot. Inst. Akad. nauk SSSR, VII, 12 (1938) 263.-7. clavatum 
141 S. Nikit. ex Zakir. Fl. i Rast. Zeravsh. 2 (1962) 413, sphalm.— Ic: 
Kuthath. Kavk. Predst. Tragop. (1957) Plate V, Fig. 16; Borissova, op. 
cit. Plate XLV, Fig. 1. 

Biennial. Root cordlike, robust, vertical; root collar covered with 
renmants of old leaves. Plants glabrous, 30-100(120) cm high, usually 
branched in lower part. Lower and middle cauline leaves linear-lanceolate 



141 

and linear, with slightly broadened semiamplexicaul base. Peduncles 
slightly thickened below capitulum, 3-6 cm thick, longer than capitula. 
Capitula cylindrical, large, 6-8 cm long, 1-1.8 cm wide. Involucral bracts 
seven or eight, rarely 12, almost two times as long as florets, as long as 
achenes or slightly longer, lanceolate-linear or linear, acuminate, 
membranous along margin in lower part; florets dark violet, raspberry- 
colored or purple. Achenes 16-50 per capitulum, straight, whitish, with 
pappus 6-7 cm long, without pappus 3.2-4 cm long; body of achene 
1.3-2 cm long, 2 mm wide, indistinctly five-angled in cross section, 
finely sulcate, finely and densely acutely tuberculate, scabrous; gradually 
narrowed into slender, bent, 1.5-2.7 2.7(3) cm long beak, clavate at 
apex, smooth and glabrous, scabrous in lower part, and with hairy annulus 
at apex; pappus 2-3.5 cm long, with isolated long hairs, dirty white, 
one and one-half to two times as long as body of achene. Flowering 
April to May; fruiting May to August. 

Mostly in the foothills, in mountains on steppe slopes, screes and 
cobbly dry slopes, sometimes in old fields and crops. — Caucasus: Eastern 
and Southern Transcaucasia, Talysh; Soviet Central Asia: Aralo-Caspian 
(Beleuty), Tien Shan (southern Kirgizia), Pamiro-Alai, mountainous 
Turkmenia. General distribution: Distribution in Iran possible. Described 
from Gaudan in Kopetdag Range. Type in Leningrad. 

Note. T. longirostris Bischof, belongs to this series, which differs 
from T. krascheninnikovii S. Nikit. in pubescence, achene with indistinct 
edges, length of the pappus, peduncles, cauline leaves, etc. 

Section 6. Tragopogon. — Peduncles not thickened below capitula 
or weakly thickened at fruiting. Capitula solitary, usually medium, on 
long straight peduncles; involucral bracts 6-11, often seven or eight; 
florets of various shades of yellow, pale violet, or purple; beak of achene 
slender, smooth, as long as body of achene or slightly shorter; pappus 
slightly shorter, as long or slightly longer than achene with beak; biennial 
or, less often, perennial plants. 

Type of section: T. pratensis L. 

Series 1. Orientales Artemcz. in Uch. Zap. Chernov Univ. I (1948) 
88, p. p. Peduncles not thickened or slightly thickened at fruiting; 
involucral bracts 7-11; florets yellow, whitish-yellow, or orange, 
sometimes pale lilac on drying. Mostly biennials. 

142 12. T. orientalis L. Sp. pi. (1753) 789; Ldb. Fl. Ross. II, 786; 

Boiss. Fl. or. Ill, 747; Kryl. H. Zap. Sib. XI, 2977; C. Rgl. in Ber. 
Schweiz. Bot. Gesellsch. 61, Map 4. — T. pratense auct. fl. ross. p. p. — 
T. pratense p. orientale Schmalh. Fl. II (1897) 146; Kyrl. Fl. Alt. Ill, 
729; Fedtsch. and Fler. Fl. Evrop. Ross. 1043.— Ic: Hegi, 111. Fl. VI, 2, 



142 

Taf. 275, f. 4; Javroka and Csapody, Iconogr. Fl. Hung. (1929) 3993. 

Biennial. Root string-like, vertical, cylindrical. Plants 12-60(90) cm 
high, glabrous, with individual flocculose scales. Stems sulcate, erect, 
simple or branched. Leaves light-colored, glaucous-green, linear or linear- 
lanceolate, acute, flat or sinuate, slightly narrowed in upper part and 
broadened near base, amplexicaul; lower leaves 10-40 cm long, 3-18(24) 
mm wide, upright, sometimes arcuate; upper leaves amplexicaul, shorter 
and wider, 3-8 cm long, 3-10 mm wide, long-acuminate, lanceolate, 
sometimes sinuate. Peduncles not thickened below capitula; capitula large, 
one each on long glabrous peduncles. Involucral bracts 8-10, lanceolate, 
2.5^ cm long, 8-12 mm wide, with greenish- white margin, much shorter 
than peripheral florets and almost as long as achenes or slightly shorter. 
Florets bright-colored, golden-yellow, or orange-yellow, up to 15 mm 
long, often whitish on drying; anthers yellow, with dark brown or almost 
black longitudinal stripes. Peripheral achenes with pappus about 3.9 cm 
long, without pappus 2.2 cm long; body of achene curved, weakly five- 
angled, scabrous or sharp-toothed-tuberculate from somewhat acute or 
subobtuse scales, sometimes shallow sulcate or smooth, almost glabrous 
in lower part, gradually narrowed into slender, about 0.9 cm-long beak, 
which is broadly inflated at apex and with hairy annulus, longer than 
body of achenes or shorter; pappus about 1.6 cm long, much shorter 
than achene, whitish. Flowering May to August (September). 

Meadows and dry slopes, forest glades, old fields and loamy soils 
in pine forests. A common, widely distributed plant. — European Part: 
Volga-Kama, Upper Volga, Upper Dnieper, Middle Dnieper, Volga-Don, 
Trans- Volga, Black Sea Region, Lower Don, Lower Volga; Western 
Siberia: Upper Tobol; Eastern Siberia: Angara-Sayans (introduced into 
Kransnoyarsk and up to Sayans foothills, near Posolsk in Baikal); Soviet 
Central Asia: Aralo-Caspian Region. General distribution: Central 
Europe. Described from western Europe. Type in London. 

Note. According to Nikitin, T. orientalis L. is a southeastern species, 
whose range comes in contact in the mountain systems of the Siberian 
water divide, with the range of T. songoricus. According to Artemczuk, 
T. orientalis L. is a Balkans-Asia Minor species. The opinion of S.A. 
Nikitin is closer to reality. T. orientalis L. is distributed in the forest- 
steppe and steppe zone of the RSFSR (T. pratensis L. is the species 
replacing it to the northwest in the forest belt). The varieties recognized 
are: T. orientalis var. revolutus Bisch. — with arcuate, sinuate leaves; 
var. laevigatus Beck. — with an indistinct sculpturing of the achenes; 
and var. tortile C.A.M. — with undulate, weakly convolute leaves. 

143 13. T. pratensis L. Sp. pi. (1753) 789; DC. Prodr. VII, 113; Ldb. 

Fl. Ross. II, 785; Schmalh. Fl. II, 146; Kryl. Fl. Zap. Sib XI, 2978; 



143 

Stank, and Tal. Opred. Vyssh. Rast. 689; C. Rgl. in Berichte Schweiz. 
Bot. Gesellsch, 65 (1955) Karte 4. — T. pratense a. vulgaris Schmalh. 
op. cit.— r. minor Fries, Novit. fl. Suec. ed. 2 (1828) 241.— (?) T. 
carinatum Gilib. Fl. Lithuan. I (1792) 183.— Ic: Fedtsch. and Fler. Fl. 
Evrop. Ross. Fig. 1066; Hegi, 111. H. VI, 2, t. 275, f. 5. 

Biennial. Root vertical, robust, cylindrical, with numerous slender 
and short lateral roots; root collar covered with fibers. Plants glabrous, 
30-90(120) cm high; stems simple or branched, straight, mostly robust, 
sometimes colored. Leaves long, linear or lanceolate-linear, basally 
semiamplexicaul, broadened and abruptly narrowed, finely long- 
acuminate, sometimes undulate along margin, almost carinate in upper 
part; middle and upper cauline leaves much shorter than lower ones. 
Peduncles long, significantly exceeding leaves, straight, glabrous, not 
thickened below capitula. Capitula solitary terminal on stem and 
branches, at flowering 2.5-3 cm long, 12-20 mm wide, elongated at 
fruiting. Involucral bracts 8-10, lanceolate or linear-lanceolate, 2.5-4 
cm long, 8-12 mm wide, long-acuminate, sometimes brown or pale in 
lower part, sometimes shorter than peripheral florets and achenes, white- 
bordered. Receptacle glabrous, initially flat, later on weakly convex, 
punctate-alveolate. Florets light yellow; anthers yellow, sometimes turning 
black at apex. Achenes with pappus 3-4 cm long, without pappus 2-2.3 
cm long, blackish or brownish-gray; body of achene slightly curved, 
sulcate, scabrous due to small subobtuse tubercles along ribs, somewhat 
thick, about 1.1 cm long, abruptly narrowed into 0.8-1 cm-long, slender 
and smooth beak as long as body of achene, not swollen at apex, and 
with hairy annulus; pappus 1-1.5 cm long, shorter than achene with 
beak, whitish. Flowering May to September. 

Meadows, meadow slopes, forest glades, borders of fields, roadsides 
and old fields. — European Part: Karelia-Lapland, Dvina-Pechora, Baltic 
Region, Ladoga-Ilmen, Upper Volga, Upper Dniester. General 
distribution: Scandinavia, central Europe, Atlantic Europe. Described 
from western Europe. Type in London. 

Note. This species replaces T. orientalis L. in the northwest. It is a 
fodder and honey-bearing plant. 

14. T. bjelorussicus Artemcz. in Tr. Inst. Bot. Khar'k. Univ. II 
(1937) 31; Klokov in Vizn. Rosl. URSR, 590; Stank, and Tal. Opred. 
Vyssh. Rast. 691; Mikhail, in Fl. BSSR, V, 177.-7. brevirostris DC. 
subsp. bjelorussicus (Artemcz.) C. Rgl. in Ser. Hort. Bot. Univ. Vytauti 
Magn. V (1937) 41; C. Rgl. in Berichte Schweiz. Bot. Gesellsch. 65 
(1955) Karte 1. — Ic: Artemcz. op. cit. Fig. 1, p. 32; C. Rgl. op. cit. 
(1937) Abb. 4 and 14. 

Biennial. Root vertical, fusiform; root collar with renmants of old 



144 

dry leaves. Plants 14-35 cm high, whitish-flocculose-tomentose, densely 
144 below capitula and near leaf base. Stems solitary or few, often branched 
from base. Basal leaves narrowly linear, 10-25 cm long, broader at 
base and flat, 2-5 mm wide, whitish-lilac below, sometimes folded, not 
exceeding fertile branches; cauline leaves wider, narrowed toward apex 
and bent, subulately acuminate, 2-6 cm long; upper leaves reduced, 2- 
4 mm wide at base, 2-6 cm long, gradually subulately acuminate. 
Peduncles below capitula not thickened; involucral bracts 7-11, at fruiting 
elongated up to (25)30-40 mm, basally 5-9 mm wide, shorter than florets, 
subulate, sometimes violet. Florets whitish-yellow, whitish on drying, 
peripheral florets more or less one and one-half times as long as 
involucre. Achenes yellowish, smooth, roundish, five-angled, 3.3-3.5 
cm long with and 1.7 cm long without pappus, about 1 mm wide, very 
shallow- and obtusely ribbed and finely squamose; beak 0.5-0.7 cm 
long, more or less one-third as long as achene, glabrous; pappus whitish- 
golden, 1.2-1.7 cm long, shorter than achenes, with short hairy annulus 
at base. Flowering June to July (September). (Plate XI, Fig. 1.) 

Coastal sands and pine forests. — European Part: Upper Dnieper 
(Pripyat and Dnieper valleys). Endemic. Described from Mozyr District. 
Type in Leningrad. 

Note. Artemczuk (op. cit.) describes var. trachycarpus Artemcz. 
with grayish cylindrical achenes having 10 scabrous ribs. 

15. T. trachycarpus S. Nikit. in Bot. Mat. Gerb. Bot. Inst. Akad. 
Nauk SSSR VII, 12 (1937) 259; Popov, Fl. Sredn. Sib. II, 72.-7. 
pratense var. griseocinerascens Genesch. in Tr. Bot. Muz. Akad. Nauk, 
XVIII (1915) 224. 

Biennial. Root cordlike, vertical, robust; root collar covered with 
remnants of previous year's leaves Plants glabrous, 50-90 cm high. Stems 
branched in upper part, strong, erect, densely leafy. Leaves not exceeding 
peduncles; lower leaves linear, cauline leaves semiamplexicaul, 10-15(26) 
mm wide, basally narrowed into filiform, long-folded lamina; upper 
leaves shorter. Peduncles 7-10 cm long at fruiting, not or very weakly 
thickened below capitulum. Capitula with mature achenes 40-50 mm 
long, 15-20 mm at flowering. Involucral bracts eight, almost as long as 
florets and achenes; young involucral bracts in lower part sericeous, 
brownish, sometimes with red dots. Peripheral florets light yellow above, 
dirty violet below (according to Ganeschin). Peripheral achenes with 
pappus about 3.5 cm long, without pappus about 2 cm long; achenes 
without beak 1 cm long, shallow-sulcate, five-angled, weakly scabrous 
along ribs or almost smooth; beak slender, smooth, about 1 cm long, 
not swollen above (without knob), with weakly developed tuft of hairs; 
pappus about 1.5 cm long; usually shorter than achene with beak, 
brownish. Flowering June to July; fruiting July to August. 



145 

145 Meadows, along shores, sometimes as weed in gardens. — Eastern 

Siberia: Dauria. Endemic. Described from Transbaikal Region (Akhur 
River). Type in Leningrad. 

Note. T. trachycarpus S. Nikit. is widespread at the eastern boundary 
of the genus. It is distinguished from T. sibiricus Ganeschin by the light 
yellow and dirty violet, not dark red-purple, color of the corolla below, 
smooth achenes (and not sulcate and weakly scabrous) with the beak 
longer than the achene, involucral bracts almost as long as florets and 
achenes, and also by the form and length of the leaves. Popov (op. cit.) 
suggested a hybrid origin for T. trachycarpus S. Nikit. (T. orientalis x 
T. sibiricus), which is doubtful, because these two species are not found 
together. 

Series 2. Turkestanici Boriss. Peduncles not or weakly thickened 
below capitula at fruiting; involucral bracts seven or eight (nine), as 
long as or slightly shorter than achenes; florets yellow, orange, or pale 
yellow, pale lilac on drying; achenes with slender beak, transition to 
body of achene abrupt. Biennial, less often annual, montane species of 
Soviet Central Asia and Altai. 

Of the species assigned to this series, T. karelinii S. Nikit. and T. 
tomentosulus Boriss., possibly belong to other groups; material of these 
species is insufficient in the collections. 

16. T. songoricus S. Nikit. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 
1, I (1933) 198; Kryl. Fl. Zap. Sib. XI, 2979. 

Biennial. Root vertical, robust; root collar covered with dry remnants 
of previous year's leaves that are basally broad. Stems rather robust, 
25-50 cm high, erect, glabrous, more or less branched from middle. 
Lower leaves narrowly linear, sometimes dropping off after flowering, 
more or less amplexicaul; middle leaves linear-lanceolate, broader at 
base, semiamplexicaul, narrowed toward apex into linear-lanceolate 
lamina; upper cauline leaves oblong-lanceolate and oblong, long- and 
filiform-acuminate, basally broadened and almost amplexicaul. Peduncles 
5-11 cm long, not thickened at fruiting; capitula 3.5-4 cm long, with 
18-40 achenes. Involucral bracts seven or eight (nine), slightly exceeding 
florets at flowering, very rarely slightly shorter than achenes, sometimes 
with dark dots at base, membranous along margin, sometimes sericeous 
below; florets yellow. Peripheral achenes with pappus 3.2-3.6 cm long, 
without pappus 1.7-1.8 cm long; body of achene 1-1.2 cm long, curved, 
finely ribbed, five-angled, finely tuberculate only along ribs, rather 
abruptly narowed into slender, smooth 0.6-0.8 cm long beak, without 
hairy annulus and thickening at apex; pappus 1.6-1.8 cm long, dirty 
white. Flowering July to August. 

In mountains on meadows and dry slopes, in feather grass-herb 



146 

146 Steppes, thickets of shrubs, moist meadows, forest edges. — Western 
Siberia: Altai (southwest); Soviet Central Asia: Balkhash Region, 
Dzhungaria-Tarbagatai, Tien Shan. General distribution: China, 
Mongolia. Described from Ketmen Range. Type in Leningrad. 

Note. T. songoricus S. Nikit. is the species replacing T. turkestanicus 
S. Nikit. ex Pavl. to the north of the Trans-IIi Alatau. It differs from T. 
turkestanicus S. Nikit. ex Pavl. by longer involucral bracts, smaller 
capitula, and a shorter beak without a hairy annulus at its apex. From 
T. orientale L., it is distinguished by an involucre that exceeds the 
florets, a shorter beak abruptly joining the broader part of the achene, 
and by the sculpture of the achenes. A food and fodder plant. 

17. T. turkestanicus S. Nikit. ex Pavl. in Byull. Mosk. Obshch. 
Isp. Prir. XLII, 2 (11 XII 1933) 136; S. Nikitin in Tr. Bot. Inst. Akad. 
Nauk SSSR, Ser. 1, I (28 XII 1933) 196; Kovalevskaja in Fl. Uzb. VI, 
433. — T. pratense auct. fl. As. Med. p. p. — T. pseudopratense M. Pop. 
in herb. — Ic: Pavlov, op. cit. Fig. 3. 

Biennial. Root robust, mostly vertical, root collar covered with dry 
remnants of previous year's leaves. Stems (25)50-100 cm high, mostly 
solitary, glabrous, erect, robust, simple, less often more or less branched 
in upper part. Lower leaves flat, linear or linear-lanceolate, 5-7 mm 
wide, like other leaves broadened at base, semiamplexicaul, acuminate 
at apex; middle leaves linear or linear-lanceolate, 3-4 mm wide, with 
wide, almost amplexicaul base, much shorter than lower leaves, abruptly 
narrowed upward into linear lamina, sometimes folded; upper cauline 
leaves reduced, oblong-lanceolate, shorter than internodes, filiform- 
acuminate, almost amplexicaul. Peduncles 5-11 cm long, at fruiting 
weakly thickened below capitula; capitula 3.5-5 cm long at fruiting. 
Involucral bracts seven or eight(nine), deltoid-lanceolate, 22-25 mm long, 
one and one-half times as long as florets, as fruiting as long as achenes 
or slightly shorter, sometimes reddish-brown and membranous along 
margin, sometimes lanate below; florets yellow. Peripheral achenes 
weakly concave, with pappus about 4 cm long, without pappus 2.2 cm 
long; body of achene about 1.2 cm long, sulcate, scabrous from fine 
tubercles along ribs, grayish, abruptly narrowed into slender, 1.1 cm- 
long beak, ribbed below, swollen at apex and with hairy annulus; pappus 
1.8 cm long, dirty-white, shorter than achene with beak. Flowering May 
to July; fruiting June to September. 

Foothills to forest and subalpine zones, on screes and stony slopes 
of ravines, in mountain meadows, at 1,000-2,600 m. — Soviet Central 
Asia: Tien Shan, Pamiro-Alai. Endemic. Possibly distributed in 
neighboring parts of Afghanistan. Described from Talass Alatau. Cotype 
in Leningrad. 



147 

Note. S.A. Nikitin described var. massarifense S. Nikit., growing in 
the Zeravshan and Turkestan ranges (op. cit. 196), on the basis of 
147 larger capitula and achenes (up to 4.7 cm long) and a longer beak on 
the achenes (up to 1.4 cm long).] 

18 T. badachschanicus Boriss. in Addenda XXVIII, 723. 

Biennial. Root robust, long, vertical; root collar covered with dry 
white sheaths of previous year's leaves. Plants glabrous; stems 15-35 
cm high, erect, branched from base. Lower leaves numerous, linear, 
0.5-1.5 cm wide, abruptly narrowed toward apex, long-acuminate, folded, 
semiamplexicaul, not exceeding peduncles, 7-20 cm long; middle cauline 
leaves much shorter than lower ones, linear to lanceolate, 
semiamplexicaul, with many veins, sometimes with narrow white border. 
Peduncles exceeding leaves, at fruiting more or less thickened below 
capitula. Capitula at flowering about 2 cm long, and about 1 cm wide, 
at fruiting 3.5-5 cm long, numerous. Involucral bracts eight(nine), 
equaling florets at flowering or slightly shorter, at fruiting equaling or 
slightly exceeding achene, green, linear-lanceolate, gradually acuminate; 
florets pale yellow, pale violet on drying. Peripheral achenes slightly 
curved, with pappus 4.3 cm long, without pappus 2.6 cm long; body of 
achene about 1.3 cm long, five-angled, finely sulcate, scabrous from 
fine tubercles, abruptly narrowed into slender, smooth about 1.2 cm 
long beak, not swollen at apex, with hairy annulus at pappus base; pappus 
about 1.7 cm long, whitish-golden, brownish or dirty white in upper 
part, shorter than achene with beak. Flowering July. (Plate IX, Fig. 1.) 

On stony steppe slopes, in grass patches along beds of 
mountain streams, at 3,200^,200 m. — Soviet Central Asia: Pamiro-Alai. 
Endemic. Described from locality of Kok-Dzhar in Pamir. Type in 
Leningrad. 

19. T. altaicus S. Nikit. and Schischk. in Bot. Mat. Gerb. Bot. Inst. 
Akad. Nauk SSSR, VII, 12 (1937) 260.-7. orientate auct. fl. alt. ex 
parte. 

Biennial. Root vertical, robust; root collar covered with remnants 
of previous year's leaves. Plants glaucous-green, glabrous, 30-60 cm 
high; stems branched from base or below middle, or simple, rather robust, 
3-5 mm thick, sulcate, sometimes colored. Leaves not exceeding stem, 
semiamplexicaul, up to 10-20 mm wide in lower part; lower leaves 
linear and lanceolate-linear, crowded or remote, long-acuminate; middle 
and upper cauline leaves shorter, abruptly narrowed, sometimes contorted 
at apex. Peduncles strong, 3-5 mm thick, weakly broadened below fertile 
capitula. Capitula with broad base, at fruiting 1.5-3 cm wide, 2-4 cm 
long. Involucral bracts 8(10), lanceolate, broader at base, acuminate and 
divergent, as long as achenes, shorter than florets by 10-20 mm; florets 



148 

orange. Peripheral achenes slightly curved, with pappus about 3 cm 
148 long, without pappus about 1.7 cm long, sulcate, obtusely ribbed, with 
alternate, finely scabrous and smooth ribs, almost without sculpture; 
body of achene 2-3 mm wide, abruptly narrowed into 5-7 mm long 
beak, not swollen at apex and with annulus of scanty hairs, very finely 
sulcate, almost smooth; pappus 1.3 cm long, shorter than achene with 
beak. Flowering and fruiting June to August. 

Mountain slopes. — Western Siberia: Altai. Endemic. Described from 
altai, from Sailyugem Mountain. Type in Leningrad 

Note. It is distinguished from T. orientalis L. by the broader capitula, 
the shape, size and sculpture of the achenes, a beak that is shorter than 
the widest part of the achene, and the color of the corolla. From T. 
pratensis L. it differs by involucral bracts that are shorter than the florets, 
corolla color, achene shape, a shorter beak, as well as by the form of 
the capitula and leaves. 

20. T. karelinii S. Nikit. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk 
SSSR, VII, 12 (1937) 237. 

Biennial or perennial. Root vertical, virgate. Plants 25-30 cm high; 
stems glabrous, finely sulcate. Leaves remote, with more or less broader 
base, 5-8(12) mm wide, semiamplexicaul, narrowed into linear-laneolate 
lamina, weakly membranous along margin. Peduncles below capitula 
not thickened, about 3 mm thick. Involucral bracts seven or eight, almost 
as long as florets and much shorter than achenes. Capitula 20-30 mm 
long at flowering; 35-50 mm long at fruiting; florets pink (dry). 
Peripheral achenes with pappus about 4.8 cm long, without pappus 2.5 
cm long; body of achene about 1.3 cm long and 1.5 mm wide, weakly 
sulcate, with scarcely noticeable scales, five-angled gradually narrowed 
into slender, about 1 . 1 cm long beak, almost as long as body of achene 
with weakly developed hairy annulus above; pappus about 2.3 cm long. 
Flowering May to June(?). 

In semidesert. — Soviet Central Asia: Balkhash Region (Lake 
Balkhash). Endemic. Described from northern Kazakhstan. Type in 
Leningrad. 

21. T. tomentosulus Boriss. in Fl. Turkm. VII (1960) 298 and in 
Addenda 381. — 71 conduplicatus auct. fl. As. Med. non S. Nikit. 

Biennial or perennial. Root virgate, weak, slightly thickened; root 
collar covered with fibrous remnants of old leaves. Stem 20-25 cm 
high, erect, somewhat slender, weakly branched, densely leafy, with 
leaves crowded at base. Leaves filiform-linear to lanceolate, appressed 
white-tomentose at base; lower leaves longer, exceeding others, reaching 
up to inflorescence, finely sulcate, finely acuminate, with somewhat 



149 




149 Plate Vffl. 

1 — Tragopogon macropogon C.A. Mey.; 2 — Ceropogon glaber L.; 
3 — Trogopogon kasachstanicus S. Nikit. 



150 

150 broader base, semiamplexicaul; upper leaves short, wider, long-acuminate, 
linear to oblong-lanceolate. Peduncles scarcely exceeding or equaling 
leaves, not thickened, 4-6 cm long, finely pubescent, tomentose below 
capitula. Capitula solitary, terminal, medium, 2-2.5 cm long at flowering, 
about 4 cm long and 1.5 cm wide in middle at fruiting. Involucral 
bracts eight, lanceolate, acuminate, pubescent at flowering, not exceeding 
florets, much shorter than achenes with pappus; florets violet. Achenes 
(immature) about 17 mm long with beak; body of achene sulcate, weakly 
scabrous above; beak about 6 nmi long, slender, weakly scabrous below, 
more or less tomentose above, almost half as long as achene; pappus 
considerably longer than achene, about 2.5 cm long, brownish, with 
unequal plumose hairs. Flowering May to June; fruiting June. 

Herb slopes of foothills up to zone of grassy steppes. — Soviet Central 
Asia: mountainous Turkmenia. General distribution: Distribution in Iran 
possible. Described from Kopetdag Range. Type in Leningrad. 

Series 3. Graciles Boriss. Peduncles not thickened; involucral bracts 
six, less often seven or eight, shorter or almost as long as florets and 
achenes; florets yellow, pale hlac or pinkish-violet on drying. Short 
perennials, branched from base, caespitose. 

22. T. gracilis D. Don in Mem. Wem. Soc. Ill (1820) 414; Wall. 
Prodr. Fl. Nep. (1824-1826) 62; DC. Prodr. VII, 115; Hook. f. Fl. Brit. 
Ind. Ill, 417; Nikitin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, 
VII, 12, 272; Rech. f. Symb. Afghan. II (1955) 191; Kitam. Fl. Afghan. 
(1960) 452.— r. pamiricus S. Nikit. in Ikonn. Opred. Rast. Pamira (1963) 
246, descr. ross. 

Perennial. Root virgate, straight. Plants 10-20(25) cm high, 
caespitose, usually branched from base, young branches flocculose; stem 
slender. Leaves remote, narrowly linear, upright, often folded, slightly 
broader at base. Peduncles below capitula not thickened. Capitula 
narrowly cylindrical at flowering, 1.8-3 cm long, about 3-3.5 cm long 
at fruiting. Involucral bracts six (less often seven or eight), shorter than 
florets and achenes; florets yellow, pale lilac on drying, exceeding 
involucral bracts. Peripheral achenes about 3.2 cm long with pappus, 
about 1.6 cm long without pappus, weakly curved; body of achene about 
1 cm long, indistinctly angular, weakly sculptured, gradually narrowed 
into slender, about 0.7 cm-long beak shorter than achene, and with tuft 
of soft hairs and apical swelling; pappus about 1.5 cm long, almost as 
long as achene with beak or shorter. Flowering July to August. 

In alpine zone. — Soviet Central Asia: Pamiro-Alai (Alai Range, 
Pamir, Shugnan). General distribution: Indo-Himalayas. Described from 
India. Type in London. 



J 



151 

151 23. T. subalpinus S. Nikit. in Bot. Mat. Gerb. Bot. Inst. Akad. 

Nauk SSSR VII, 12 (1937) 271. 

Perennial or biennial. Root vertical, thick; root collar covered with 
remnants of previous year's leaves. Plants relatively small, 10-25 cm 
high, branched usually from base, caespitose. Stems slender, simple or 
branched, densely leafy. Leaves semiamplexicaul, broader in lower part, 
5-10 mm wide, linear-lanceolate or linear (lower leaves), sometimes 
undulate, more or less incurved. Peduncles below capitula not thickened, 
slender, 1.5-3 nmi wide. Capitula 1.5-2 cm long at flowering; 3-3.5 
cm long at fruiting; involucral bracts (seven) eight, acuminate, lanceolate, 
almost as long as florets and achenes; florets pink- violet. Achenes with 
pappus 2.3 cm and without pappus 1.5 cm long; body of achene 0.9 cm 
long, almost smooth, weakly scabrous in upper part or entirely smooth, 
weakly sulcate, rather abruptly narrowed into about 0.6 cm-long smooth 
beak, sUghtly swollen at apex and without hairy annulus; pappus about 
1.3 cm long, slightly shorter than achene with beak or almost as long. 
Flowering June to July; fruiting July to August. 

Steppe slopes in alpine zone. — Soviet Central Asia: Dzhungaria- 
Tarbagatai, Tien Shan. General distribution: Dzhungaria-Kashgaria. 
Described from Dzhungarian Alatau. Type in Leningrad. 

Note. It differs from T. gracilis D. Don by its involucre of eight 
bracts almost as long as the florets, pink-violet (dry) florets, shape of 
the leaves and achenes, as well as by its range. 

Series 4. Purpurascentes Boriss. Peduncles not or weakly thickened; 
florets purple or pinkish-violet; involucral bracts 8-12, almost as long 
as florets and achenes or slightly longer; plants biennial. 

24. T. sibiricus Ganesch. in Tr. Bot. Muz. Akad. Nauk XIII (1915) 
225; XVI (1916) 129; Kryl. Fl. Zap. Sib. XI, 2973; Popov, R. Sr. Sib. 
II, 72.— Ic: Ganeschin, op. cit., Plate XV. 

Biennial. Root thickened above middle, vertical, long. Plant glabrous, 
light green, 45-110 cm high; stem erect, smooth, sometimes in upper 
part, as also leaves, reddish-violet. Leaves linear or broadly linear, 
broader at base, 4-8(13) mm wide, semiamplexicaul; lower leaves 20- 
35 cm long; middle cauline and upper leaves strongly ovately broadened 
at base and abruptly narrowed in upper part, 4-12(16) cm long. Peduncles 
weakly thickened below capitula, covered with ferruginous and black 
hairs. Capitula 15-20 mm long, at fruiting 30-50 mm long, solitary 
terminal on stem and branches. Involucral bracts 8(10), 1.5-3 cm long, 
almost as long as achenes; florets dark red or purple; peripheral florets 
two-thirds as long as involucre. Achenes with pappus about 4 cm long, 
without pappus about 2.3 cm long; body of achene smooth, roundish; 



152 

152 without sculpture, abruptly narrowed into smooth and slender 6.5-14 
mm-long beak, slightly swollen at apex and with hairy annulus; pappus 
whitish, about 1.7 cm long, almost as long as achene with beak. 
Flowering June to July. (Plate VII, Fig. 3). 

Meadows, river valleys, forest glades, open woodland^. — Western 
Siberia: Ob', Irtysh, Altai; Eastern Siberia: Angara-Sayans. Endemic. 
Described from Siberia. Type in Leningrad. 

25. T. cretaceus S. Nikit. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk 
SSSR VII, 12 (1937), 446; Fl. Yugo-Vost. VI, 446; Stank, and Tal. 
Opred. Vyssh. Rast. (1949) 688; C. Rgl. in Berichte Schweiz. Bot. 
Gesellsch. 65 (1955) Karte 2. 

Biennial. Root vertical. Plants 15-35 cm high; stem slender, glabrous. 
Lower leaves narrowly linear, 2.5-5 mm wide, almost filiform at ends, 
often folded; middle leaves broadened, semiamplexicaul, (3)4-6 mm 
wide, with linear-lanceolate lamina, narrowed into filiform cusp; upper 
leaves reduced. Peduncles sometimes slightly thickened below capitula, 
4-6 mm thick. Capitula narrowly cylindrical at flowering, 7-12(15) mm 
wide (dry), 2.5-3.5 cm long, up to 5.5 cm at fruiting; involucral bracts 
8-12, filiform-acuminate, somewhat exceeding florets and achenes; florets 
pinkish-violet or purple; outer achenes almost straight, five-angled, 
squamose-scabrous, indistinctly sculptured in lower part, gradually 
narrowed into slender beak longer than achene; pappus dirty brown, 
much shorter than achene with beak. Flowering May. 

On outcrops and chalky slopes. — European Part: Lower Volga (near 
Volsk). Endemic. Described from chalky slopes in vicinity of Volsk. 
Type in Leningrad. 

Note. It is distinguished from T. porrifolius mainly by the narrow, 
conduplicate leaves, smaller capitula, involucre slightly exceeding the 
florets and achenes, and sculpture of the achenes. 

Section 7. Tuberosi (Kuthath.) Boiss. comb. nova. — Ser. Tuberosi 
Kuthath. Kavk. Predst. Tragop. (1957) 25. — Peduncles slightly thickened 
or not below capitula; capitula medium, 2-A cm long, solitary, on straight 
peduncles; involucral bracts eight, as long as florets or shorter, almost 
as long as outer achenes or shorter; florets yellow and pale yellow, 
sometimes somewhat lilac with brownish veins on drying; achenes 
narrowed into somewhat slender, straight beak, almost as long as 
brownish-squamose body of achene; perennial plants with tuberous roots. 

Type of section: T. tuberosus C. Koch. 

26. T. tuberosus C. Koch in Linnaea XXII (1849) 662; Grossh. Fl. 
Kavk. IV, 231; Kuthath. Kavk. Predst. Tragop. (1957) 31; Kuthath. in 



153 

153 Fl. Azerb. VIII, 506. — T. pusillum acut. non M.B.: Boiss. Fl. or. Ill 
(1875) 753, p. p.— Ic: Kuthath. in Fl. Gruzii VIII (1952) Fig. 428; 
Kuthath. op. cit. (1957) Plate III, Fig. 12, Map 5.— Exs.: PI. or. exs. 
No. 99. 

Perennial. Root mostly vertical, swollen at some depth as globose 
or cylindrical, sometimes elongated tuber; root collar sometimes covered 
with sheaths of dead leaves. Plants 20-40(45) cm high, glabrous or 
flocculose; stems erect. Basal leaves linear, long, numerous; cauline 
leaves 3-10 cm long, strongly broadened at base, five- to seven-veined, 
gradually narrowed, almost amplexicaul, sometimes weakly pubescent. 
Peduncles weakly thickened below capitula. Capitula large, 2-4 cm long 
at flowering, 4-5 cm long at fruiting; florets bright yellow, with four 
violet stripes on lower side, pale violet on drying. Involucral bracts 
eight, as long as ligulate florets or shorter, lanceolate, almost as long as 
achenes with pappus at fruiting. Peripheral achenes with pappus 3.7^.3 
cm long, without pappus 1.6-2.4 cm long; body of achene 1.1-1.4 cm 
long, tuberculate-squamose along ribs and deeply sulcate, gradually 
narrowed into ribbed, glabrous, about 1 cm-long beak, shorter than body 
of achene, with hairy annulus at apex; pappus 1.7-1.9 cm long, white 
or grayish, shorter than achene with beak; inner achenes narrower, 
indistinctly ribbed. Flowering April to May; fruiting May to June. 

Dry stony slopes, among scrub up to upper mountain zone. — 
Caucasus: Ciscaucasia, Dagestan, Eastern Transcaucasia, Southern 
Transcaucasia. Endemic. Described from Georgia. Type was in Berlin. 

27. T. pusillus M.B. Fl. taur.-cauc. Ill (1819) 521; DC. Prodr. VII, 
115; Ldb. Fl. Ross. II, 789; Boiss. Fl. or. Ill, 753, p. p.; Grossh. Fl. 
Kavk. IV, 232; Borissova in Fl. Turkm. VI, 296— T. pratense p. 
eriophorum DC. Prodr. VII (1837) 113.— Ic: Kuthath. in Fl. Gruzii 
VIII, Fig. 430; Kuthath. Kavk. Predst. Tragop. (1957) Plate III, Fig. 13, 
Map 5. 

Perennial. Root long, vertical, slender, swollen at some depth, as 
mostly oblong tuber, deep in soil; root collar covered with hairs or 
sheaths of previous year's leaves. Plants short, 5-15(25) cm high, loosely 
flocculose in lower part, glabrescent. Leaves numerous; basal leaves 
narrowly linear, slightly broadened at base, semiamplexicaul, densely 
pubescent, one- to three-veined, erect or slightly twisted and coiled at 
apex, furrowed, sometimes pubescent along margin; cauline leaves wider 
at base and shorter, oblong-lanceolate, long-acuminate. Peduncles one- 
headed, slightly exceeding leaves or as long, not thickened below 
inflorescence. Capitula 2-3 cm long. Involucral bracts eight, narrowly 
lanceolate, as long as florets or shorter; florets pale yellow, weakly lilac 
on drying with brownish veins in lower part. Achenes with pappus about 



154 

3 cm long, without pappus 1.6 cm, about 1 cm long excluding beak, 
154 about 1 mm wide in body of achene, scabrous, shallow-sulcate, gradually 
narrowed into slender, about 0.6 cm-long, rugose beak, sulcate and 
without hairy annulus at apex; pappus about 1.3 cm long, shorter than 
achene with apex; pappus about 1.3 cm long, shorter than achene with 
beak. Flowering April to May; fruiting May to June. 

Dry slopes of hills, from desert to alpine zone. — Caucasus: Dagestan, 
Eastern and Southern Trancaucasia, Talysh; Soviet Central Asia: 
mountainous Turkmenia (Greater Balkhans Range). General distribution: 
Iran. Described from Azerbaidzhan and from eastern Georgia. Type in 
Leningrad. 

Section 8. Profundisulcati Kuthath. Kavk. Predst. Tragop. (1957) 
49, p. p. — Peduncles not thickened below capitula or slightly thickened; 
capitula large; involucral bracts often 5, less often 10-12, very rarely 5, 
longer or shorter than florets; florets yellow; body of achene with five 
deep furrows and five prominent, sometimes winged squamose ribs, 
abruptly narrowed into slender beak; pappus not longer than achenes; 
plants biennial. 

Type (lectotype) of section: T. armeniacus Kuthath. 

Series 1. Subobconico-pedunculati Kuthath. Kavk. Predst. Tragop. 
(1957) 51 p. p. — Peduncles not or weakly thickened below capitula; 
involucral bracts 8, less often 10-12, shorter than florets; florets yellow; 
beak as long as body of achene; pappus mostly shorter than or as long 
as achenes. 

28. T. meskheticus Kuthath. in Zam. po Sist. i Geograf. Rast. Tbil. 
17 (1953) 34; Fl. Gruzii VIII, 627; Kuthath. Kavk. Predst. Tragop. (1957) 
51.— Ic: Kuthath. op. cit. (1957) Plate VII, Fig. 23 (incorrect), Map 9. 

Perennial. Root vertical, cylindrical, long; root collar without old 
leaves. Plants glaucous or flocculose and spinescent. Stems numerous, 
branched, whitish, densely leafy up to apex. Basal leaves broadly linear, 
long, glaucous; cauline leaves gradually broadened at base, long- 
acuminate; uppermost leaves broadly lanceolate, short-acuminate. 
Capitula rather small. Peduncles not thickened below capitula or weakly 
thickened at fruiting; florets yellow or pale yellow; involucral bracts 
eight, lanceolate, as long as or slightly longer than achenes with pappus, 
recurved on fruiting. Peripheral achenes flat, white, deeply furrowed, 
acutely tuberculate-squamose along ribs, abruptly narrowed into beak, 
longer than body of achene; achenes shallow-sulcate, finely tuberculate, 
with shorter beak; beak with white hairy annulus at apex; pappus shorter 
than achene with beak. Flowering July; fruiting August. 



155 

155 On saline soils in lower mountain zone. — Caucasus: Eastern 

Transcaucasia (Meskhetia). Endemic. Described from Meskhetia. Type 
in Tbilisi. Cotype in Leningrad. 

Note. Description of achenes taken from Kuthatheladze (1957), 
where, however, the illustration (Plate VII, Fig. 23) does not correspond 
with the characters of the species described in the text. 

29. T. ketzkhovelii Kuthath. in Zam. po Sist. i Geogr. Rast. Tbil. 
16 (1951) 96; Kuthath. Fl. Gruzii VIII, 628; Kavk. Predst. Tragop. (1957) 
52. — T. reticulatus var. latifolius Sosn. ex Kuthath. op. cit. (1951) 98. — 
Ic: Kuthath. op. cit. (1951) 97, Fig. 1; op. cit. (1957) Plate VIII, Fig. 
27, Map 9. 

Biennial. Root vertical, cylindrical; root collar often covered wtih 
remnants of dead basal leaves. Plants almost glabrous, 30-55 cm high, 
with erect, thick stems, simple or branched from base, angular in cross 
section. Basal leaves long, almost half as long as plant, linear, not 
broadened at base; cauline leaves basally broadened, gradually narrowed, 
acuminate. Capitula large, solitary. Peduncles long, far exceeding leaves, 
not thickened below capitula; involucral bracts eight, lanceolate, slightly 
pubescent at base, almost as long as achenes with pappus or slightly 
shorter; florets yellow, as long as involucral bracts or shghtly longer. 
Peripheral achenes with pappus 5.5 cm long, without pappus 2.5 cm 
long; body of achene 1.3 cm long, about 2,5 mm wide, grayish, deeply 
sulcate, winged, with five prominent winged ribs covered with upright, 
short, denticulate spinules, abruptly narrowed into straight, somewhat 
slender grayish beak almost as long as body of achene, about 1.2 cm 
long, scabrous and finely sulcate below, slightly swollen at apex and 
with hairy annulus; pappus about 2.5 cm long, with isolated nonplumose 
hairs, up to 3.2 cm long, light golden-yellow; inner achenes almost 
indistinguishable from peripheral. Flowering June to August. 

Mountain slopes of the middle mountain zone in more or less 
xerophytic habitats. — Caucasus: Southern Transcaucasia (Armenian 
uplands and Dzhavakhetia). Endemic. Described from Dzhavakhetia in 
vicinity of village of Kumurdo. Type in Tbilisi. 

30. T. plantagineus Boiss. and Huet in Boiss. Diagn. pi. or. ser. II, 
3 (1856-1859) 91; Kuthath. Kavk. Predst. Tragop. (1957) 53.— T. 
buphtalmoides a. latifolius Boiss. Fl. or. Ill, 751; Grossh. Fl. Kavk. IV, 
233.— r. brevifolius C. Koch in Linnaea XXIII (1850) 662.— Ic: 
Kuthath. op. cit. Plate VIII, Fig. 26, Map 9. 

Biennial. Plants up to 70 cm high; root vertical; cylindrical; root 
collar covered with few hairs. Stems cylindrical, simple, sometimes 
branched above, with straight branches. Basal leaves oblong-broadly 



156 

lanceolate, 8-10 cm long and (11)20-30 cm wide; cauline leaves reduced, 
156 very short at forks, short-acuminate, recurved. Peduncles slightly 
thickened below capitula. Capitula narrow in beginning of flowering, 
later broadening; florets yellow. Involucral bracts eight, shorter than 
achenes with pappus. Peripheral achenes whitish-grayish, large, with five 
deep furrows and five prominent winged ribs, abruptly narrowed into 
beak almost as long as body of achene; inner achenes with shallow 
furrows, acutely squamose, gradually narrowed into somewhat thick beak; 
pappus dirty gray, with hairy annulus at base. Flowering V-VI; fruiting 
VI-VII. 

Mountain slopes. — Caucasus: Southern Transcaucasia (Armenia). 
General distribution: Asia Minor. Described from vicinity of Erzurum. 
Type in Geneva. 

31. T. buphtalmoides (DC.) Boiss. Fl. or. Ill (1875) 750; Grossh. 
Fl. Kavk. IV, 233; C. Rgl. in Berichte Schweiz. Hot. Gesellsch. 65 
(1955) Karte 3; Kuthath. Kavk. Predst. Tragop. (1957) 54.— T. 
buphtalmoides p. stenophyllum Boiss. op. cit. 751. — T. persicum Boiss. 
Diagn. pi. or. ser. I, 7 (1846) 4; Boiss. Fl. or. Ill, 751; in syn. Grossh. 
op. cit. — Scorzonera buphtalmoides DC. Prodr. VII (1838) 121. — Ic: 
Kuthath. op. cit. Plate VIII, Fig. 28, a (incorrect). Map 9. 

Perennial or biennial. Root vertical, cylindrical; root collar often 
covered with previous year's leaves. Plants 40-70 cm high, flocculose; 
stems rather densely leafy, erect, simple or branched from base. Basal 
leaves linear or lanceolate, up to 25 cm long, 7-8 nun wide, usually 
seven-veined, often withering during flowering; cauline leaves linear, 
shorter, gradually narrowed toward apex; upper cauline leaves reduced, 
2-2.5 cm long, broadened at base, semiamplexicaul, short-acuminate, 
with distinct longitudinal veins. Peduncles at fruiting slightly thickened 
below capitula, smooth or pubescent. Capitula large; florets yellow. 
Involucral bracts 8-10(12), often tomentose near base, shorter than 
achenes and florets, sometimes incurved at fruiting. Peripheral achenes 
with beak about 5.3 cm long, without pappus about 2.3 cm long; body 
of achene about 1.5 cm long, deeply five-furrowed and obtusely ribbed 
or winged, dentate-squamose along ribs, abruptly narrowed into somewhat 
thick, finely sulcate, about 0.6 cm-long beak, swollen at apex and with 
hairy annulus; pappus about 3 cm long, lustrous, golden or grayish- 
golden; inner achenes almost smooth and terete, shallow-ribbed, gradually 
narrowed into beak. Flowering May to June; fruiting June to July. 

Dry stony and cobbly slopes, in maple-juniper woodlands in middle 
and upper mountain zones. — Caucasus: Southern Transcaucasia, Talysh. 
General distribution: Balkans- Asia Minor, Armenia and Kurdistan, Iran. 
Described from northern Syria. Type in Paris. 



157 

Note. A polymoq)hic plant; vars. stenophyllus Boiss. and humilis 
Boiss. are distinguished on the basis of leaf width and plant height. 

157 32. T. tasch-kala Kuthath. in Zam. po Sist. i Geogr. Rast. Tbil. 17 

(1953) 38; Kuthath. Kavk. Predst. Tragop. 2 (1957) 55.— Ic: Kuthath. 
op. cit. (1957) Plate VII, Fig. 24, Map 9. 

Biennial. Root vertical, cylindrical; root collar often covered with 
dry leaves. Plants flocculose, 30-50 cm high. Stems cylindrical, whitish, 
branched from base or middle, densely leafy, with one capitulum each 
at branch apices. Basal leaves linear, weakly broadened toward base, 
gradually narrowed toward apex, up to 15 cm long, not longer than 
stem; cauline leaves dense, numerous, reduced, 4-6 mm long, lanceolate, 
broadened at base to 7-10 mm, semiamplexicaul, gradually narrowed 
toward apex, peduncles longer than leaves, slightly thickened at fruiting, 
fistular, glabrous or tomentose at base of capitula. Capitula large, up to 
4.5 cm long at fruiting; florets yellow, much longer than involucral 
bracts. Involucral bracts 12, lanceolate, whitish, involute (dry), shorter 
than florets and achenes with pappus. Peripheral achenes about 4.5 cm 
long, excluding pappus about 2.3 cm long, slightly curved; body of 
achene excluding beak about 1.5 cm long, sulcate-furrowed, with five 
prominent acutely squamose winged ribs and with five small furrows, 
with erect scales, nonsquamose below, rather abruptly narrowed into 
0.8 cm-long, slender, sulcate beak, swollen at apex and with hairy 
annulus; pappus yellowish, about 2 cm long; inner achenes slightly longer 
than peripheral ones. Flowering July. 

In subalpine zone, at about 1,750 m. — Caucasus: Southern 
Transcaucasia. Endemic. Described from Armenia (Karabari, Tashkala). 
Type in Tbilisi; isotype in Leningrad. 

33. T. karjaginii Kuthath. Kavk. Predst. Tragop. (1957) 57; Fl. 
Azerb. VIII, 511.— Ic: Kuthath. op. cit. Plate IX, Fig. 29 (incorrect). 
Map 10. 

Biennial. Root vertical; root collar often covered with previous year's 
leaves. Plants glabrous, glaucous. Stems branched from base, whitish, 
up to 60 cm high. Basal leaves longer than cauline leaves; cauline leaves 
broadened at base, finely and long-acuminate, semiamplexicaul, remote. 
Peduncle weakly thickened below capitula, fistular. Capitula large. 
Involucre bracts eight, shorter than achene with pappus; florets unknown. 
Peripheral achenes about 3.5 cm long, slender, abruptly narrowed into 
beak, rather deeply narrow-sulcate, with 10 ribs, white squamose along 
ribs, curved, convex in middle or below, grayish; inner achenes 
ferruginous, shallow-sulcate, flater and squamose, almost smooth, 
especially in lower part, with beak as long as or slightly longer than 



158 

body of achene, hairy (?) at apex below pappus; pappus about 1.7 cm 
long, shorter than achene with beak. Flowering May to June; fruiting 
June to July. 
158 Stony slopes of lower mountain zone. — Caucasus: Eastern 

Transcaucasia (Azerbaidzhan). Endemic. Described from the vicinity of 
Dzhebrail. Type in Baku. 

Series 2. Latifolii Kuthath. Kavk. Predst. Tragop. (1957) 44.— 
Peduncles not thickened below capitula; involucral bracts 8-10 or 10- 
12, shorter than ligulate florets; outer achenes with five more or less 
deep furrows and five ribs, gradually narrowed into beak, shorter than 
achenes; florets yellow. 

34. T. latifolius Boiss. Diagn. pi. or. ser. I (1844) 23; Boiss. Fl. or. 
Ill, 750; Sosn. in Tr. Azerb. Otd. Zakavk. Fil. Akad. Nauk SSSR, I, 48; 
Grossh. Fl. Kavk. IV, 232; Kuthath. Kavk. Predst. Tragop. (1957) 44, 
Map 8; Fl. Azerb. VIII, 509. 

Biennial. Root straight, long, slender or thick, but not tuberous. 
Plants 50-60 cm high; stems erect, robust, angular, simple, or, mainly 
in upper part, weakly branched, densely leafy, finely sulcate, glabrous, 
flocculose below; intemodes short. Leaves oblong or broadly lanceolate, 
2-4 cm wide, abruptly narrowed toward apex, semiamplexicaul near 
base, entire and weakly undulate, 8-1 2- veined; upper leaves shorter, 
lanceolate, glabrous or appressed hairy. Peduncles not thickened below 
capitula, considerably exceeding leafy part of stem. Capitula solitary, 
rather large, 3-4 cm wide; florets yellow, exceeding involucre; involucral 
bracts 10-12, wide, lanceolate, long-acuminate, shorter than outer florets, 
flocculose at base at flowering. Outer achenes with five more or less 
deep furrows, squamose-tuberculate, gradually narrowed into beak much 
shorter than achenes; pappus whitish or dirty white, shorter than achenes 
with beak. Flowering May to June; fruiting June to August. 

Subalpine and alpine meadows and forest edges at 2,000-2,400 m, 
sometimes in crops. — Caucasus: Eastern and Southern Transcaucasia. 
General distribution: Balkans-Asia Minor. Described from Asia Minor. 
Type in Geneva. 

Note. Figure 20 in Plate VI in the work of Kuthatheladze (1957) is 
not correct. 

35. T. acanthocarpus Boiss. Diagn. pi. or. I, 11 (1849) 45; Boiss. 
Fl. or. Ill, 751, p. p.; Grossh. Fl. Kavk. IV, 232; Kuthath. Kavk. Predst. 
Tragop. 48; Kuthath. in Fl. Azerb. VIII, 509.— Ic: Kuthath. op. cit. 
(1957) Plate VI, 19, Map 8. 

Biennial. Root robust, vertical; root collar covered with fibrous 
sheaths of previous year's leaves. Plants glabrous or slightly flocculose. 



159 




161 Plate IX. 

1 — Tragopogon badachschanicus Boriss.; 2 — T. sabulosus Krasch. and S. Nikit. 



160 

up to 60 cm high; stem slender, erect, cylindrical, simple or weakly 
branched above, whitish, finely sulcate, leafy, with short intemodes. 
Basal leaves narrow, long, linear, finely and long-acuminate, usually 
folded, glabrous, withering at flowering; cauline leaves shorter, expanded 
159 in lower part, up to 10-13 mm, amplexicaul, subulately acuminate, 
usually five-veined. Peduncles not expanded below capitula, glabrous or 
flocculose. Capitula rather large, about 2.5 cm wide and 3 cm long. 
Involucral bracts 8-10, lanceolate, two-thirds as long as peripheral florets, 
about 2 cm long, erect, broadened toward base; florets yellow. Peripheral 
achenes 3.5^.5 cm long, excluding pappus, 2-2.5 cm long, 1.5 nmi 
wide, sharply five-angled, along margin acerose and verrucose or finely 
squamose, shallow-sulcate, with more or less distinct, distant, acute, 
erect, curved spinules along ribs; body of achenes gradually narrowed 
into about 1 cm-long, sulcate beak, somewhat slender, swollen at apex, 
and with hairy annulus, less than half as long as achene; pappus about 
1.7-2 cm long, dirty white or grayish, slightly shorter or almost as long 
as achene with beak. Flowering June to July; fruiting July to September. 
Dry slopes in subalpine and alpine zones, at 1,900-2,400 m. — 
Caucasus: Eastern Transcaucasia (Nakhichevan), Southern Transcaucasia. 
Talysh. General distribution: Iran. Described from Tochal Mountain in 
vicinity of Tehran. Type in Geneva; isotype in Leningrad. 

36. T. idae Kuthath. Kavk. Predst. Tragop. (1957) 46.— Ic: Kuthath. 
op.cit. Plate V, Fig. 17 (incorrect). Map 8. 

Biennial. Root vertical, cylindrical, thick, but not tuberous. Plants 
glaucous-green, up to 50 cm high; root collar often with few fibrous 
remnants of previous year's leaves. Stems erect, branched almost from 
base or middle, glabrous, cylindrical, leafy. Cauline leaves 2-5 cm long, 
1.5 cm wide, lanceolate, briefly narrowed at apex, roundish or weakly 
acuminate. Peduncles below capitula slightly expanded or not, glabrous. 
Capitula large. Involucral bracts eight, narrowly lanceolate, white- 
membranous along margin, shorter than florets and achenes with pappus; 
florets yellow. Peripheral achenes with beak up to 3 cm long, with five 
shallow furrows and five white, squamose, obtuse ribs; beak as long as 
body of achenes, clavately thickened at apex, with hairy annulus; pappus 
yellowish, lustrous, slightly shorter than achene with beak. Flowering 
August. 

Mountain meadows. — Caucasus: Southern Transcaucasia (Armenia, 
Daralagyoz District). Endemic. Described from Armenia. Type in Tbilisi. 

Note. It differs from T. acanthocarpus Boiss. by the shape and size 
of its leaves, structure of the outer achenes, ratio of beak length to broadest 
part of the achene, and general appearance. 

37. T. makaschwUii Kuthath. in Zam. po Sist. i Geogr. Rast. Tbil. 



161 

17 (1953) 33; Kuthath. in Fl. Gruzii VIII (1952) 628; Kavk. Predst. Tragop. 
(1957) 47, Map 8. — T. elatior auct. non Stev.: Grossh. Opred. Rast. Kavk. 
(1949) 506.— Ic: Kuthath. op. cit. (1953) Fig. 3. 
160 Biennial. Root vertical, cylindrical; root collar covered with fibrous 

remnants of dead leaves. Stem solitary, cylindrical, erect, sparsely leafy, 
with indistinct nodes, finely sulcate. Leaves lustrous, dark green, 
semiamplexicaul; basal and lower cauline leaves almost similar, linear or 
broadly linear, 25-30 cm long, about 1.5 cm wide; uppermost leaves 
reduced, 2-3.5 cm long, expanded at base, lanceolate, gradually attenuate 
and long-acuminate. Peduncles below capitula slightly or practically not 
thickened. Capitula solitary, large, 3.5 cm long, about 3.5 cm wide; florets 
yellow, with light blue stripes on drying. Involucral bracts 10, lanceolate, 
glabrous, about 3 cm long, shorter than florets. Achenes unknown. 
Flowering August. 

In crop fields of the lower mountain zone. — Caucasus: Eastern 
Transcaucasia (Akhalkalaki District) and Southern. Transcaucasia. 
Endemic. Described from Georgia. Type in Tbilisi; isotype in Leningrad. 

Series 3. Armeniaci Boriss. Peduncles not or slightly thickened below 
capitula at fruiting; involucral bracts five, shorter than achenes; outer 
achenes sulcate, with five prominent winged ribs, abruptly terminating 
in thick sulcate beak, with hairy annulus at apex; pappus yellowish. 

38. T. armeniacus Kuthath. in Zam. po Sist. i Geogr. Rast. Tbil. 
17 (1953) 36; Kuthath. Kavk. Predst. Tragop. (1957) 56, Map 9.— Ic: 
Kuthath. op. cit. (1953) Fig. 5; op. cit. (1957) Plate VII, Fig. 25. 

Root cylindrical, vertical; root collar without previous year's leaves. 
Plants glabrous, up to 45 cm high. Stems cylindrical, erect, simple or 
branched above, with single capitulum at branch apices. Basal leaves 
narrowly linear, folded; cauline leaves shorter, expanded at base, 
semiamplexicaul, folded. Peduncles below capitula not or weakly 
thickened at fruiting, glabrous or covered with very short, appressed 
yellowish hairs. Capitula at fruiting large. Involucral bracts five, 
lanceolate, gradually long-acuminate, shorter than achene with pappus, 
less often as long; florets purple(?). Peripheral achenes large, about 4 
cm long with pappus, without pappus 1.9 cm; body of achenes 1.2 cm 
long, 4 mm wide, sulcate, with five prominent winged ribs with sharp 
dentate margin and five indistinct, sharply dentate ribs in between, 
abruptly narrowed into thick, sulcate, about 0.6 cm-long beak, slightly 
swollen at apex and with hairy annulus; pappus about 2.3 cm long, 
yellowish. Flowering June; fruiting July. 
162 In subalpine zone, at 2,000 m. — Caucasus: Southern Transcaucasia 

(former Artik District). Described from Armenia. Type in Tbilisi; isotype 
in Leningrad. 



162 

Section 9. Bessera Boiss. sect. nov. in Addenda XXVIII, 724. — 
Peduncles not thickened below capitula, scarcely exceeding leaves; 
capitula mostly small and medium; involucral bracts (seven) eight or 8- 
12, shorter than florets and achenes; florets yellow or pale yellow; 
achenes slender, flat, smooth or weakly scabrous, narrowed into indistinct, 
1 .5-3 mm long beak; pappus slightly shorter than achene; plants perennial 
(less often biennial) flocculose, especially at flowering, sometimes 
caespitose, simple or branched, densely leafy; root collar covered with 
fibrous remnants of old leaves; leaves narrow to filiform. On sands. 

Type of section: T. lithuanicus (DC.) Boriss. 

Note. Is distinguished from the species of section Brevirostres 
Kuthath. by the caespitose habit, number of involucral bracts (8-12 and 
not five to eight), and shape of the achenes; achenes slender, flat and 
enlongated, mostly smooth, with an indistinct beak and a pappus shorter 
than the achene; in the species of section Brevirostres; achenes shallow- 
sulcate, tuberculate-scabrous with a distinct, short beak. 

Series 1. Lithuanici Boriss. ser. nov. Perennial prostrate plants with 
numerous creeping vegetative branches, forming turf; involucral bracts 
eight; capitula rather small; florets light yellow; achenes about 2.3 cm 
long, almost smooth, with indistinct, about 3 mm-long, somewhat thick 
beak; pappus shorter than achene. 

39. T. lithuanicus (DC.) Boriss. comb. nova. — T. brevirostre 5. 
Uthuanicum DC. Prodr. VII (1838) 114; Ldb. Fl. Ross. II, 1 (1844) 
788. — T. floccosus subsp. lithuanicus (Bess.) C. Rgl. in Ser. Hort. Bot. 
Univ. Vyt. Magni V (1937) 43; C. Rgl. in Berichte Schweiz. Bot. 
Gesellsch. 65 (1955) Map 1. 

Perennial. Root virgate, robust; rhizome branched, bearing numerous 
prostrate vegetative branches, forming dense turf. Stems short, fertile, 
5-10 cm long, densely white-flocculose-arachnoid, glabrous at fruiting, 
simple or branched, densely leafy. Basal leaves filiform and narrowly 
linear, 3-8 cm long, 0.5-1 mm wide, sparsely flocculose-arachnoid; 
middle and upper leaves abruptly acuminate from oblong-ovate or 
lanceolate base to narrowly linear apex, sometimes deflexed, 7-30 mm 
long, 1-3.5 mm wide at base, flocculose-arachnoid, especially at leaf 
base. Involucral bracts eight, 2.5-4 mm wide, 12-22 mm long at base, 
linear-lanceolate, thin-acuminate, densely flocculose-arachnoid at 
flowering. Peduncles slightly exceeding leaves, glabrous. Florets light 
163 yellow, peripheral florets slightly exceeding involucre. Achenes with 
pappus about 2.3 cm long, without pappus 13-15 mm long, fusiform, 
body of achene 10-13 mm long, smooth or weakly scabrous and finely 



163 

sulcate, in upper part gradually narrowed into less distinct, about 3 mm- 
long beak; pappus shorter than achene, 10-14 mm long. Flowering July 
to August. 

On sands. — European Part: Baltic Region (Lithuania). Described 
from Lithuania (Rukhcha). Type in Kiev; isotype in Leningrad. 

Note. A unique plant, not collected again by anyone; apparently a 
rare relict. 

Series 2. Heterospermi Boriss. — Perennial(?) branched plants; 
involucral bracts (7)8-12; capitula 2.5-3.5 cm long; florets yellow or 
light yellow, exceeding involucral bracts; achenes with pappus 2.5-3.5 
cm long, thin, smooth or finely squamose, scabrous along ribs, gradually 
and smoothly narrowed toward apex; beak not developed; pappus shorter 
than achene. 

40. T. gorskianus Rchb. Ic. Fl. Germ. XIV (1860) 19, t. 43; 
Artemcz. in Tr. Inst. Bot. khar'k. Univ. II, 31; Stank, and Tal. Opred. 
Vyssh. Rast. 690; C. Rgl. in Scr. Hort. Bot. Univ. Vyt. Magni V (1937) 
47; Berichte Schweiz. Bot. Gesellsch. 65 (1955) Map 1. — T. floccosus 
subsp. heterospermus var. gorskii C. Rgl. 1. c. (1955) 252. 

Perennial(?). Root long, fusiform, vertical; root collar covered with 
remnants of old leaves. Stem branched, ascending. Basal leaves narrower 
than cauline leaves, 2-5 mm wide. Involucral bracts longer than florets; 
florets yellow, shorter than involucre. Peripheral achenes smooth, with 
beak as long as body of achene; pappus pale brown. Flowering June(?). 

On sands. — European Part: Baltic Region (vicinity of Druskenikai 
in Lithuania, Neman River). Endemic. Described from Lithuania. Type 
unknown. 

Note. I did not have at my disposal material for this species. 

It is distinguished from T. floccosus Waldst. and Kit., with which it 
is synonymized, by size of stem, ratio of beak and achene length, and 
length of achene. From T. bjelorussicus Artemcz. it differs by the 
relationship of the length of the beak and the achene and by yellow 
florets that are shorter than the involucral bracts; from T. orientalis it 
differs by having smooth achenes. 

41. T. tanaiticus Artemcz. in Tr. Inst. Bot. Khar'k. Univ. II (1937) 
51; Klok. in Vizn. Rosl. 591. — T. ruthenicus auct. non Bess,: S. Nikit. 
in Majevski, FI. (1933) 705; Nikit. in Izv. Glavn. Bot. Sada SSSR, 
XXIX, 5-6, 487, ex. p. — T. ruthenicus subsp. tanaiticus (Artemcz.) C. 
Rgl. Scr. Hort. Bot. Univ. Vyt. Magni V (1937) 44; C. Rgl. in Berichte 
Schweiz. Bot. Gesellsch. 65 (1955) Map 2. — Ic: Artemcz. op. cit. Fig. 
7; C. Rgl. op. cit. Abb. 8, 19. 



164 

Perennial(?). Root robust, vertical; root collar covered with remnants 
of old leaves. Plant 15-75 cm high, more or less densely flocculose- 
164 arachnoid, especially below capitula. Stems solitary or few, branched, 
rather densely leafy. Upper leaves reduced, 1.5-5 cm long, not convolute, 
expanded at base up to (4)8-12 mm, abruptly narrowed into linear, 2- 
5(8) mm wide lamina, acute; cauline leaves 8-10(16) mm wide at base, 
toward apex sometimes curved or involute, gradually narrowed; basal 
leaves narrower than cauline leaves, narrowly linear, expanded at base, 
whitish-lilac, folded, acute. Involucral bracts (seven) eight, shorter than 
florets, narrowly lanceolate, 18-25 mm long, at fruiting (4)8-10 mm 
wide, narrowly membranous along margin, usually pubescent, sometimes 
alternating with less pubescent bracts, glabrescent. Capitula medium; 
peduncles below capitula not thickened; florets yellow. Peripheral achenes 
five-angled, thin, 2.5 cm long with and 1.3-1.5 cm long without pappus, 
sulcate, finely squamose, scabrous along ribs, slightly narrowed toward 
apex, beak not developed; pappus 9-10 mm long, grayish, shorter than 
achene, with hairy annulus near base; inner achenes smooth, with pappus 
about 2.8 cm long. Flowering June to August. 

Sands in steppe and forest-steppe zone in the middle reaches of the 
Don and Volga. — European Part: Middle Dnieper (Kharkov), Volga- 
Don, Lower Don, Lower Volga. Endemic. Described from Voronezh 
Region (village of Berez-nyagi). Type in Leningrad. 

Note. It is distinguished from T. ruthenicus (C.A.M.) Bess, by 
shorter, more compact leafy stems, narrower leaves, thinner, curved, 
five-angled, and less scabrous achenes, and a longer pappus. From T. 
floccosm Waldst. and Kit. it differs by the absence of the beak. 

42. T. heterospermus Schweigg. in Koenigsb. Arch. Naturw. I 
(1812) 229; Hagen, Chloris Borussica (1818) 145; ibid (1819) 288; S. 
Nikit. in Izv. Glavn. Bot. Sada SSSR XXIX, 5-6, 490; Artemcz. in Tr. 
Inst. Bot. Khar'k. Univ. II, 33. — T. brevirostre 5. floccosum (Waldst. 
and Kit.) Schmalh. Fl. II (1897) 147. — T.floccosus subsp. heterospermus 
(Schweigg.) C. Rgl. in Scr. Hort. Bot. Univ. Vyt. Magni V (1937) 42; 
C. Rgl. in Berichte Schweiz. Bot. Gesellsch. 65 (1955) Karte 1.— Ic.:. 
Rchb. Ic. Fl. Germ. XIX, 1 (1860) 19, t. 41.— Exs.: GRF Nos. 332, 
3799. 

PerenniaK?) or biennial. Root vertical, long, cylindrical; root collar 
covered with fibrous remnants of old leaves. Plants 20-50 cm high, 
lanate-tomentose, sometimes mainly in lower parts; stem usually branched 
from base, densely leafy. Leaves not exceeding stem; lower leaves linear, 
expanded at base, often folded; cauline leaves amplexicaul, expanded at 
base, 4-10 nmi wide, 3-10 cm long, narrowed upward, not deflexed, 
subulate; upper leaves reduced, 4—5 mm wide at base, 10-15 mm long, 



165 

gradually subulately acuminate, sometimes more or less incurvate. 
Peduncles not thickened below capitula. Capitula numerous, solitary 
terminal on branches, 3^ cm long. Involucral bracts 8-12, at flowering 
165 23-30 mm long, shorter than florets and achenes, lanceolate, acuminate, 
with elongated cusp; florets pale yellow, exceeding involucral bracts. 
Peripheral achenes with pappus 2.5-3.5 cm long, without pappus 
1.3-1.6 cm, about 1 nmi wide, sulcate, five-ribbed, finely tuberculate or 
almost smooth, almost without or with weakly developed, about 2-3 
mm-long beak, one-sixth to one-fifth as long as body of achene; pappus 
about 1.5 cm long, yellowish, with short hairy annulus at base. Flowering 
May to August. (Plate VII, Fig. 1.) 

Sand dunes. — European Part: Baltic Region (near Riga, in Lithuania, 
Kaliningrad Region). Endemic. Described from Baltic Region. Topotype 
in Leningrad. 

Note. It differs from T. floccosus Waldst. and Kit. by plant size 
and by having more densely lanate-tomentose pubescence, numerous 
flat, usually less reduced cauline leaves, larger capitula and achenes, 
and a separate range. From T. bjelorussicus Artemcz. it differs by having 
broader achenes, shorter involucral bracts and achenes with a shorter 
beak. C. Regel (op. cit.) recognizes var. maritimus C. Rgl. on the sandy 
shores of the Baltic Sea and var. heidenreichii Abromeit {Fl. Ost-West- 
preussen (1903) 466) in the Kaliningrad Region, which differ by having 
achenes with a distinct beak, an almost complete absence of pubescence, 
green, glabrous stems, and a height of about 30 cm. 

Section 10. Brevirostres Kuthath. Kavk. Predst. Tragop. (1957) 12, 
p. p. — Peduncles not thickened below capitula; capitula small to medium, 
on erect, long peduncles; involucral bracts (five) six to eight, shorter 
than ligulate florets; florets yellow, sometimes pale lilac with violet 
veins on drying; achenes shallow-sulcate, tuberculate-squamose or 
smooth, distinctly narrowed into short beak or beak indistinct; pappus 
almost as long as achene with beak. Plants perennial and biennial, not 
caespitose; root collar covered with fibers or old dead leaves. 

Type of section: T. dasyrynchus Artemcz. 

Series 1. Dasyrhynchiformes Artemcz. in Uch. Zap. Chernov. Univ. 
I (1948) 89, p. p.; Kuthath. Kavk. Predst. Tragop. (1957) 22, p. p.— 
Florets yellow; involucral bracts mostly eight, less often five, shorter 
than florets; achenes short, with short beak, mostly with hairy annulus 
at apex or glabrous; capitula 1.5-2.5 cm long. Plants biennial 
predominantly of steppe zone. 

43. T. dasyrhynchus Artemcz. in Tr. Inst. Bot. Khar'k. Univ. II 
(1937) 42; Klok. in Vizn. Rosl. URSR, 590; Grossh. Opred. Rast. Kavk. 



166 

506; C. Rgl. in Berichte Schweiz. Bot. Gesellsch. 65 (1955) 252, Karte 
2.—T. brevirostre auct. nonnul. non DC: Boiss. Fl. or. Ill (1875) 752 
and others; Schmalh. Fl. II, 147, ex parte. — T. elatior auct. nonnul. 
166 non Stev.: Grossh. Opred. Rast. Kavk. 506, p. p. — Ic: Artemcz. op. cit. 
Fig. 4; Kuthath. Kavk. Predst. Tragop. (1957) Plate II, Fig. 8, Map 2. 

Biennial. Roots relatively slender, easily pulled out. Plants 25-130 
cm high, glabrous or weakly pubescent, or flocculose-arachnoid- 
tomentose near leaf base and below capitula; root collar usually covered 
with hairs. Cauline leaves 5-16(35) mm, upper ovate-lanceolate, semi- 
amplexicaul, wider, more reduced at base, narrowed upward in flat lamina 
with undulate margin, recurved at apex; basal leaves linear, 5-13 mm 
wide, often folded. Involucral bracts (seven) eight, shorter than florets. 
Peduncles not thickened below capitula. Florets yellow; anthers blackish. 
Peripheral achenes about 2.7 cm long with pappus, shallow-sulcate, 
acutely squamose-tuberculate near base, gradually narrowed into scabrous, 
somewhat thick, 1-5 mm-long beak, one-fifth to half as long as achene; 
pappus about 1.7 cm long, almost as long as achene with beak, whitish, 
easily detached, with hairy annulus at base. Flowering May to August. 

Dry steppe slopes, dry meadows in steppe and forest-steppe zone, 
scrubs and on river sands, in mountains on dry and stony slopes up to 
lower zone. — European Part: Black Sea Region, Crimea, Lower Don; 
Caucausus: Ciscaucasia (western). Endemic. Described from former Don 
Region. Type in Kiev. 

Note. According to Artemczuk, T. dasyrhynchus Artemcz. is a 
xerophytic, primarily steppe species closely related to T. ruthenicus (C. 
A.M.) Bess., T. donetzicus Artemcz. and T. borystenicus Artemcz., 
forming a series of vicariant species. I am referring T. borystenicus to 
series Stepposi Boriss., and T. ruthenicus and T. donetzicus to section 
Dubjanskya Boriss., which encompasses the psammophilous, beakless 
species. 

44. T. volgensis S. Nikit. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk 
SSSR, VII, 12 (1937) 261; Nikit. in Fl. Yugo-Vost. VI, 444.-7. 
brevirostre var. volgense S. Nikit. in Majevski, Fl. 6th ed, (1933) 706. — 
T. brevirostris subsp. volgensis C. Rgl. in Scr. Hort. Bot. Univ. Vyt. 
Magni V (1937) 40; C. Rgl. in Berichte Schweiz. Bot. Gesellsch. 65 
(1955) Karte 1.— Ic: Rgl. Ibid. Abb. 3, 13. 

Biennial. Root vertical, thickened in upper part; root collar covered 
with fibrous remnants of previous year's leaves. Plants light green, 20- 
60 cm high; stems few or solitary, simple or branched in lower part. 
Lower leaves numerous, as also other leaves, not exceeding stems, linear, 
herbaceous, without prominent veins, sometimes with whitish stripe in 
middle; and upper cauline leaves slightly expanded at base. 



167 

semiamplexicaul, 3-8(10) mm wide, linear or linear-lanceolate. Peduncles 
not thickened below capitula, Capitula elongated at fruiting, narrow, 
0.8-1.5 cm wide, 18-25(30) mm long. Involucral bracts five to eight, 
shorter or almost as long as peripheral florets, shorter than achenes, 
linear-lanceolate, acuminate; florets yellow. Peripheral achenes with 
167 pappus about 2.8 cm long, excluding pappus 1.0-1.3(1.4) cm long, 
indistinctly obtusely angular, shallow-sulcate, finely tuberculate along 
ribs, gradually narrowed into about 0.4 cm-long, sulcate beak, not swollen 
at apex and with hairy annulus; pappus about 1.2 cm long, isolated 
pappus hairs up to 1.5 cm long. Flowering May to August; fruiting July 
to September. (Plate X, Fig. 1.) 

Meadows and sands. — European Part: Volga-Don, Trans-Volga 
Region, Lower Volga; Soviet Central Asia: Aralo-Caspian Region. 
Endemic. Described from floodplain meadows of Volga near Astrakhan. 
Type in Leningrad. 

Note. The florets were described from specimens collected in the 
Volga meadows near Syzran. The species differs from T. podolicus Bess, 
by the elongated capitula, number of involucral bracts (five to eight and 
not eight), size and sculpture of the achenes, and the relationship of the 
length of the achene, pappus and beak, 

45. T. podolicus (Bess. ex. DC.) Artemcz. in Tr. Inst. Bot. Khar'k. 
Univ. II (1937) 34; Klok. in Vizn. Rosl. URSR, 590.-7. podolicus 
Bess, ex DC. Prodr. VII (1838) 114, pro syn. — T. podolicum auct. nonn.: 
S. Nikit. in Fl. Yugo-Vost. VI, 444, p. p. — T. brevirostre DC. y. 
podolicum DC. 1. c; Ldb. Fl. Ross. II, 788; Schmalh. Fl. II, 147 ex 
p. — T. brevirostris subsp. podolicus var. pratensis C. Rgl. in Scr. Hort. 
Bot. Univ. Vyt. Magni V (1937) 39. — T. brevirostris subsp. podolicus 
C. Rgl. in Berichte Schweiz. Bot. Gesellsch. 65 (1955) Karte 1.— Ic: 
Artemcz. op. cit. Fig. 2; C. Rgl. op. cit. (1955) Abb. 1, 12. 

Biennial. Root vertical, fusiform; root collar covered with remnants 
of old leaves. Plant glabrous or grayish, more or less weakly flocculose- 
tomentose, 25-90 cm high, strongly branched from middle; stems solitary 
or sometimes numerous, usually erect. Lower leaves linear, 10-25(35) 
cm long, 3-8(14) mm wide, usually with prominent veins; middle cauline 
leaves expanded at base, amplexicaul to middle, gradually narrowed 
upward, acuminate, flat or sometimes folded and pubescent, with obtuse 
callose thickening; upper leaves short, relatively broad, lanceolate-linear, 
finely and long-acuminate, sometimes apically contorted. Peduncles not 
thickened. Capitula numerous, at fruiting 20-30 mm long, in panicles at 
branch apices. Involucral bracts eight, lanceolate, acuminate, 15-20 mm 
long, shorter than florets and achenes; florets light yellow, peripheral 
florets slightly exceeding involucral bracts. Peripheral achenes smooth. 



168 

indistinctly obtusely ribbed, without sculpture, indistinctly squamose- 
scabrous at base, less often with indistinct sculpture; better developed 
achenes, acicular, with pappus about 1.8 cm long, gradually narrowed 
into beak, achenes 1.2 cm long without pappus, about 1 mm wide, with 
short 3^ mm-long beak; pappus easily detached, shorter than achenes, 
about 0.8 cm long, with soft hairy annulus at base and slightly thickened. 
Flowering May to August. 

Found infrequently along forest edges, in forest-steppe and steppe 
zones, dry meadows, on roadsides, less often in crops and on clayey- 
168 sandy substrates; in south, exclusively in meadows. — European Part: 
Upper Dnieper, Middle Dnieper, Volga-Don, Bessarabia, Black Sea 
Region, Lower Don. General distribution: Possibly, in Central Europe. 
Described from Podolia. Type in Kiev; isotype in Leningrad. 

Note. Var. floccosiformis Artemcz. (op. cit. 38) is recognized; entire 
plant lanate-arachnoid, with basal leaves up to 3 mm wide, expanded at 
the base; cauline leaves amplexicaul, expanded at the base, undulate 
along margin, annulately or spirally twisted upward; peduncles glabrous; 
achenes with a short beak. On steppe slopes and meadows of Dnieper. 

46. T. elatior Stev. in Bull. Soc. Nat. Mosc. XXIX (1956) 407; 
Boiss. Fl. or. Ill, 749; Artemcz. in Tr. Inst. Bot. Khar'k. Univ. II (1937) 
41; C. Rgl. in Scr. Hort. Bot. Univ. Vyt. Magni V, 44; C. Rgl. in 
Berichte Schweiz. Bot. Gesellsch. 65, Karte 3. — T. brevirostre auct. non 
DC: Velen. Fl. Bulg. (1891) 367; Schmalh. Fl. II, 146. 

Biennial. Root relatively slender, easily pulled out. Plants 90-125 
cm high, glabrous; stem erect, solitary, branched. Basal leaves 5-7 cm 
wide, oblong-lanceolate, more or less long-petiolate, acuminate, undulate- 
involute along margin; cauline leaves expanded at base, reduced, 
narrowed into lanceolate lamina, acuminate; upper leaves also expanded 
at base, up to 5 mm wide, subulately pointed gradually, about 10 mm 
long. Peduncles not thickened below capitula. Capitula at fruiting up to 
6 mm wide near base, at flowering 2-2.5 cm long; involucral bracts 
eight, up to 21 mm long, acute, lanceolate; florets yellow; peripheral 
florets usually longer than involucral bracts or as long. Peripheral achenes 
with pappus about 3 cm long, without pappus about 1.4 cm long, 10- 
angled, curved, sulcate-scabrous, gradually narrowed into thick, short, 
3-4 mm-long, sulcate beak; pappus grayish, 11-15 mm long, with 
scarcely hairy annulus at base. Flowering May to June (September). 

Grassy areas of lower mountain zone. — European Part: Crimea. 
General distribution: Balkans-Asia Minor (Bulgaria), Iran. Described from 
Laspi. Type in Helsinki. 

Note. The report of Grossheim (Opred. Rast. Kavk. 506) about the 
occurrence of this species along the Black Sea Coast (from Anapa to 



169 

Tuapse) should be- attributed to T. dasyrhynchus Artemcz.; T. elatior 
Stev. differs from this latter species by larger size, thicker achenes with 
a short, thick beak, and wide basal leaves. 

Series 2. Graminifolii Kuthath. Kavk. Predst. Tragop. (1957) 12.— 
Series Filifolii Kuthath. Ibid. 19, p. p.— Florets yellow or orange-yellow, 
sometimes with violet veins on drying; involucral bracts shorter than 
florets, 6-10; achene with beak much shorter than its body, or almost 
beakless; capitula small, less often medium, up to 3 cm long; plants 
perennial. 

169 47. T. graminifolius DC. Prodr. VII (1838) 114; Ldb. Fl. Ross. II, 

2, 788; Boiss. Fl. or. Ill, 752; Schmalh. Fl. II, 146; Grossh. Fl. Kavk. 
IV, 233; C. Rgl. in Berichte Schweiz. Bot. Gesellsch. 65, Karte 1; Kitam. 
Fl. Afgh. 452; Kuthath. in Fl. Azerb. VIII, 501. — T. caucasicum auct. 
non Bess.: Fisch. and Mey. ex DC. Prodr. VII (1838) 114 pro syn.— T. 
talyschense Boiss. and Buhse in Nouv. Mem. Soc. Nat. Mosc. XII (1860) 
137. — T. orientalis auct. fl. Cauc, non L. — Ic: Boiss. and Buhse op. 
cit. t. 9; Borissova in Fl. Turkm. VII, Plate XLIV, Fig. 2; Kuthath. in 
Fl. Azerb. VIII, Plate VII. 

Perennial. Root vertical, fusiform. Plants almost glabrous or, less 
often flocculose-lanate, 20-90 cm high, with numerous stems originating 
from base; root collar covered with fibers of old leaves. Stems simple 
or branched, with ascending or erect branches. Leaves graminaceous, 
green, flat, with five veins, linear or linear-lanceolate to lanceolate, 4- 
8 mm wide, weakly recurved and acuminate; upper leaves reduced, often 
expanded at base. Peduncles exceeding leaves, not thickened below 
capitula and slightly hairy. Capitula small or medium, few-flowered; 
involucral bracts eight, at fruiting 4-8 mm wide, shorter than achenes 
and slightly shorter than florets; florets pale yellow, whitish on drying, 
exceeding involucre by one-third. Peripheral achenes terete, shallow- 
sulcate, 10-ribbed, white-tuberculate-squamose or sharply acicular- 
tuberculate, 2.5-3 cm long with pappus, about 1.1 cm long without 
pappus, 1.5 mm wide, almost smooth near base; body of achene about 
0.8 cm long, gradually narrowed into finely sulcate, 3-8 mm-long beak 
slightly shorter or almost as long as achene, with hairy annulus and 
swollen at apex; pappus about 1.1 cm long, almost as long as achene 
with beak. Flowering April to June; fruiting June to September. 

Meadows, scrubs, up to middle mountain zone. — Caucasus: 
Ciscaucasia, Dagestan, Eastern, Western, and Southern Transcaucasia, 
Talysh; Soviet Central Aisa: mountainous Turkmenia. General 
distribution: Asia Minor (former Kars Region), Iran. Described from 
Imeretia. Type in Paris. 



170 

48. T. brevirostris DC. Prodr. VII (1838) 114 excl. var.; Ldb. FI. 
Ross. II, 2, 788, p. p.; Boiss. Fl. or. Ill, 799, p. p.; Schmalh. Fl. II, 146, 
p. p.; Grossh. Fl. Kavk. IV, 231; Fedtsch. and Fl. Fl. Evrop. Ross. 
1043, p. p.; Artemcz. in Tr. Inst. Bot. Khar'k. Univ. II, 27, p. p.; Kuthath. 
Kavk. Predst. Tragop. (1957) 15; Kuthath. in Fl. Azerb. VIII, 502.-7. 
brevirostris var. genuinus Schmalh. op. cit. p. p. — T. brevirostris subsp. 
genuinus C. Rgl. in Scr. Hort. Bot. Univ. Vyt. Magni V (1937) 39, p. 
p.; C. Rgl. in Berichte Schweiz. Bot. Gesellsch. 65, Karte 1, p. p. — T. 
caucasicus Bess, ex DC. 1. c. pro syn., non Fisch. and Mey. nee. Szov. — 
Ic: Kuthath. op. cit. Plate 1, Fig. 2, Map 3. 

Perennial. Plants glabrous, grayish-green, sometimes arachnoid 
or flocculose until flowering at stem base and below capitula; root collar 
170 sparsely covered with fibers or not. Stems often numerous, less often 
solitary, 30-60(90) cm high, 1.5-3 mm thick, usually branched above 
base, with erect virgate branches bearing solitary capitula. Basal leaves 
long, slightly expanded at base, linear or linear-lanceolate, 2-4 mm wide; 
cauline leaves linear or linear-lanceolate, 1.5-4 mm wide at base, 
subulately acuminate; upper leaves reduced, 1-1.5 mm wide at base, 
folded into subulately narrowed lamina, strongly recurved at apex, up 
to 21 cm long. Peduncles oblong, glabrous, not thickened below capitula, 
sparsely leafy. Capitula small, 5-7 mm wide at base; involucral bracts 
eight, almost as long as florets or shorter, bracts lanceolate-linear, 20- 
25 mm long, 3^ mm wide at base, sometimes 5-7 mm wide at fruiting, 
acuminate; florets yellow, slightly exceeding involucre, violet on drying. 
Peripheral achenes 2.5-2.8 cm long, about 1 mm wide, acutely dentate- 
tuberculate-squamose at base, with 12 ribs covered by whitish scales, 
shallow-sulcate, gradually narrowed into glabrous, smooth, white beak 
clavately thickened below pappus; beak 3-6 mm long, achenes without 
beak about 1.3 cm long; pappus longer than achene with beak, white, 
10-15 mm long, with hairy annulus at base. Flowering April to May; 
fruiting June to August. 

Grows in upland steppes, sometimes on sands and limestones, on 
coastal sands and rocks. — Caucasus: Ciscaucasia, Eastern and Wastem 
Transcaucasia, Dagestan. Endemic. Described from the Caucasus on the 
basis of plants raised by Besser from seeds supplied by Steven. Isotype 
in Leningrad; type in Paris. 

Note. The beak of the achene is almost absent in the plants collected 
in Dagestan. 

49. T. serotinus Sosn. in Kuthath. in Zam. po Sist. i Geogr. Rast. 
Tbil. 16 (1951) 98; Kuthath. in Fl. Gruzii VIII, 635; Kavk. Predst. Tragop. 
(1957) 18; in Fl. Azerb. VIII, 502.— Ic: Kuthath., op cit. (1951) Fig. 2; 
op cit. (1957) Plate 1, Fig. 3, Map 2; MakashviH, Fl. Okr. Tbil. II, 223. 



171 

Perennial. Plants 35-50 cm high, sparsely flocculose; root collar 
covered with fibers and previous year's dry leaves; stems numerous, 
branched, densely leafy, ascending. Leaves narrowly linear; basal leaves 
longer, cauline leaves reduced, gradually acuminate, usually slightly 
falcate, folded. Peduncles exceeding leaves, slender, not thickened below 
capitula, densely pubescent. Capitula numerous, small; florets pale 
yellow, as long as or slightly longer than involucre; involucral bracts 
six to eight, lanceolate. Peripheral achenes about 2.5 cm long, excluding 
pappus about 1.4 cm long, shallow-sulcate, squamose-tuberculate, 10- 
ribbed, obtusely angular, with two rows of solitary indistinct scales, 
almost smooth at base, narrowed at apex into about 0.5 cm-long beak, 
almost half as long as body of achene (about 0.9 cm long), glabrous or 
171 slightly hairy below pappus; pappus about 1.2 cm long, shorter than 
achene with beak. Flowering April to September; fruiting August to 
November (up to late auturmi). 

Stony dry slopes and hills, coastal sands, in lower and middle 
mountain zones, at 900-1,000 m. — Caucasus: Eastern Transcaucasia 
(vicinity of Tbilisi, Kartlia, Meskhetia). Endemic. Described from the 
vicinity of Tbilisi. Type in Tbilisi. 

Note. It differs from T. graminifolius DC. by its very slender and 
strongly branched stems, narrower leaves, densely pubescent peduncles, 
small capitula, paler yellow florets, general appearance, and time of 
flowering. 

50. T. filifolius Rehm. ex Boiss. Fl. or. Ill (1875) 754; Grossh. Fl. 
Kavk. IV, 232; Kuthath. in Fl. Azerb. VIII, 504.-7. filifolium var. 
macrorhizum Somm. and Lev. in Tr. Peterb. Bot. Sada XVI (1901) 
291.— Ic: Kuthath. in Fl. Gruzii VIII, Fig. 432; Kavk. Predst. Tragop. 
(1957) 19, Plate 1, Fig. 4 (incorrect). 

Perennial. Root vertical, slender or thick, but not tuberous; root 
collar densely covered with fibers. Stems less numerous, usually two or 
three, ascending, not branched at base, 10-30 cm high. Basal and lower 
cauline leaves narrow, filiform, about 2.5 mm wide, sometimes to 
fibriform, 5-7 cm long, recurved, lanate-flocculose near sheaths, 
numerous, other leaves glabrous; cauline leaves expanded at base up to 
4 nmi. Peduncles much exceeding basal leaves, tomentose below capitula. 
Capitula about 2.5 cm long; involucral bracts 8-10, erect, about 10 mm 
long, narrow, acuminate, tomentose at base, shorter than florets; ligulate 
florets orange-yellow with purple stripes, exceeding involucre by one- 
fourth. Outer achenes about 2.4 cm long with pappus, without pappus 
about 1 cm long; body of achene weakly ribbed and scabrous along 
ribs, mainly in upper part, about 7 mm long, gradually narrowed into 
sulcate, short, 0.3-0.4 cm-long beak slightly swollen at apex and with 



172 

hairy annulus; pappus about 1 cm long, some pappus bristles longer and 
scabrous (at apex); inner achenes almost smooth. Flowering May to 
July; fruiting June to August. 

In rock crevices and meadows of alpine and subalpine zones. — 
Caucasus: Ciscaucasia, Eastern and Southern Transcaucasia, Dagestan. 
Endemic. Described from Daryal ravine. Type in Krakow, holotype in 
Tbilisi. 

51. T. colchicus Alb. Probr. fl. Colch. (1895) 151; Grossh. Fl. Kavk. 
IV, 232; Kuthath. in Fl. Gruzii VIII, 631; Kavk. Predst. Tragop. (1957) 
20.— Ic: Kuthath. op. cit. (1957) Plate II, Fig. 6 (incorrect). Map 2. 

Perennial. Root thick, cylindrical, but not tuberous; root collar 
densely covered with fibers. Stems 20-30 cm high, numerous, erect. 
172 Basal leaves numerous, narrowly linear, slightly narrowed toward base, 
much shorter than peduncles, up to 20 cm long, 1-1.5 nmi wide; upper 
cauline leaves semiamplexicaul, filiform-acuminate; upper leaves 4-5 
cm long. Peduncles exceeding leaves, erect, not thickened below capitula. 
Capitula medium, 2-3 cm long, tomentose at base; involucral bracts 
seven or eight, lanceolate, finely acuminate, expanded at base to 0.5 
cm, 2-2.5 cm long; florets yellow or orange-yellow, two times as long 
as involucre. Achenes with pappus about 2 cm long, without pappus 
about 1 cm long, 1 mm wide, smooth, obtusely sulcate, not scabrous, 
almost without beak; pappus about 0.9 cm long, white or gayish white. 
Flowering May to July. 

Meadows in alpine and subalpine zones, on limestones and rocks, 
at 2,000-2,350 m. — Caucasus: Western Transcaucasia. Endemic. 
Described from Mingrelia. Type in Tbilisi. 

Series 3. Ste pposi Boriss. Involucral bracts seven or eight, as long 
as florets or shorter; capitula small, florets yellow; achenes small, beak 
of achene not developed or indistinct; pappus almost as long as achene; 
biennial meadow-steppe plants, also growing on sands. 

52. T. stepposus (S. Nikit.) Stank, in Stank and Tal. Opred. Vyssh. 
Rast. (1949) 691. — T. podolicum subsp. stepposum S. Nikit. in Bot. Mat. 
Gerb. Bot. Inst. Akad. Nauk SSSR, VII 12 (1937) 261; in H. Yugo- 
Vost. VI, 444. — T. podolicum auct. fl. ross. et fl. turk. p. p. non DC, 
nee Artemcz.: Kryl. Fl. Zap. Sib. XI, 2975. — T. brevirostris subsp. 
podolicus var. stepposus C. Rgl. in Scr. Hort. Bot. Univ. Vyt. Magni V 
(1937) 40.— T. brevirostre auct. fl. ross.: Kryl. Fl. Alt. Ill (1904) 730, 
non DC. — T. brevirostre var. angustifolium S. Nikit. in Majevski, Fl. 
6th ed. (1933) 706.— Ic: C. Rgl. op. cit. Abb. 2. 

Biennial. Root vertical, cylindrical; root collar covered with remnants 
of old dry leaves. Plants grayish-green, 30-90 cm high. Stems slender, 
numerous, branched from middle, initially more or less arachnoid, 
flocculose, glabrescent. Lower leaves narrowly linear, 1-3(6) mm wide, 



173 

usually folded; middle cauline leaves flat, slightly expanded at base, 
narrow to filiform upward; upper leaves lanceolate to oblong-lanceolate, 
4-14 mm wide. Peduncles not thickened below capitula. Capitula 
relatively small, at fruiting 17-25(30) mm long; involucral bracts seven 
or eight, 15-22 mm long, shorter than florets and achenes or almost as 
long as latter; florets light yellow. Peripheral achenes about 2.3 cm 
long including pappus, excluding pappus 1 cm long, 1.5 mm wide, 
slightly curved, sulcate, with distinct sculpture, usually with undeveloped 
scales, gradually narrowed toward apex, almost without beak, without 
swelling (nob); pappus slightly longer than achene, about 1.3 cm long; 
inner achenes about 2.6 cm long with pappus, 1.3 cm long without 
pappus, straight, about 1 mm wide, weakly scabrous, without beak, 
pappus about 1.5 cm long. Flowering June to July. 
173 Steppe and forest-steppe zones, on steppe slopes, alkaline steppes; 

desert-steppe zone. — European Part: Volga-Don, Lower Volga, Trans- 
Volga Region; Western Siberia: Upper Tobol, Irtysh, Altai; Soviet Central 
Asia: Aralo-Caspian, Balkhash Region. Endemic. Described from vicinity 
of Lake Zaisan. Type in Leningrad. 

Note. It differs from T. podolicus Bess, by the narrowly linear leaves, 
often folded and straight (and not involute), small capitula, beakless 
achenes without thickening at the tip, sculpture and size of the achene 
and pappus, as well as the range. It differs from T. scoparius S. Nikit. 
by the narrowly linear, straight leaves, slender stems, small heads, less 
deeply sulcate achenes, and range. 

53. T. ucrainicus Artemcz. in Tr. Inst. Bot. Khar'k. Univ. II (1937) 
38; Klok. in Vizn. Rosl. URSR, 590; C. Rgl. in Berichte Schweiz. Bot. 
Gesellsch. 65 (1955) Karte 2. — T. canus auct. non Willd.: Bess. Enum. 
pi. Volh. (1822) 30.— (?) T. villosum auct. non L.: Pall. Reise II (1778) 
332, 464, p. p.; Ill, 537, 654, 682; DC. Prodr. VII, 115.-7. floccosus 
auct. fl. ucr. non Waldst. and Kit. — Ic: Artemcz. op. cit. Fig. 3; C. 
Rgl. op. cit. Abb. 6, 17. 

Biennial. Root long, vertical; root collar covered with dry old leaves 
or fibers. Plants 30-80 cm high, more or less densely pubescent, 
canescent mainly on young branches and below capitula, arachnoid- 
tomentose. Stems solitary, often pyramidally branched from base. Basal 
leaves narrower than cauline, linear, 2-3 mm wide, flat or folded, 
expanded at base, pale violet, with long lamina; cauline leaves reduced, 
expanded at base to 6-7 mm, with somewhat long lamina, undulate, 
acuminate, contorted or spirally twisted; upper cauline leaves as also 
middle, spirally twisted or annular, reduced, 10-50 mm long, 4-8 nrni 
wide at base. Peduncles not thickened below capitula; involucral bracts 
(seven) eight, 15-23 mm long at fruiting, 5-10 mm wide, basally 
tomentose; florets yellow, exceeding involucral bracts, less often as long. 



174 

Achenes with pappus about 2.3 cm long, without pappus 1.3 cm long, 
smooth or weakly scabrous above or throughout, gradually narrowed 
into short glabrous, somewhat thick, about 3-4 mm-long beak less often 
half as long as achene; pappus short, shorter than achene, about 1 cm 
long, with erect hairs and short hairy annulus at base. Flowering May 
to September. 

Sands in forest-steppe and steppe zone, pine forest sands, wastelands, 
and old sandy fields. — European Part: Upper Dnieper, Middle Dnieper, 
Volga-Don (along Donets), Lower Don. Endemic. Described from 
Kharkov Region. Type in Kiev. 

Note. Artemczuk recognizes var. lasiocarpus Artemcz. (op. cit.) with 
a scabrous achene and smooth beak, one-fourth as long as the achene; 
f. strioantherus Artemcz. — with an anther tube having five black stripes 
at the tip; and f. flaviantherus Artemcz. — with a pale yellow anther tube. 

174 54. T. daghestanicus (Artemcz.) Kuthath. in Zam. po Sist. i Geogr. 

Rast. Tbil. 17 (1953) 42; C. Rgl. in Berichte Schweiz. Bot. Gesellsch. 
65 (1955) Karte 2; Kuthath. Kavk. Predst. Tragop. (1957) 2A.—T. 
dasyrhynchus subsp. daghestanicus Artemcz. in Tr. Inst. Bot. Khar'k. 
Univ. II (1937) 44; C. Rgl. op. cit. 251.— T. floccosus auct. fl. cauc. 
non Waldst. and Kit. — T. ruthenicus auct. fl. cauc. non Bess. — Ic: 
Kuthath. op. cit. (1953) Fig. 8; op. cit. (1957) Fig. 9, Map 2. 

Biennial. Root robust, hard, cylindrical; root collar usually covered 
with fibers. Plants up to 60 cm high, entirely densely grayish-flocculose- 
arachnoid, especially in upper part. Stems solitary, branched in upper 
part, less often from base. Basal leaves linear, long, sometimes folded; 
lower cauline leaves linear but expanded at base; upper leaves reduced, 
gradually narrowed from broad base into rather long, thin acuminate 
apex. Peduncles not thickened below capitula. Florets yellow; involucral 
bracts eight, lanceolate, about 25 mm long, shorter than achene with 
pappus; achenes including pappus about 3.2 cm long, excluding pappus 
about 1.4 cm long, without beak, shallow-sulcate above, with two rows 
of scales, rest smooth; pappus about 1.8 cm long, longer than achene, 
with annulus of short hairs at base. Flowering May to June. 

Semidesert steppes among hummocky sand dunes, along the coast 
of the Caspian Sea and in foothill steppes. — European Part: (?) Trans- 
Volga Region, Lower Don, Lower Volga (Ergeni); Caucasus: Ciscaucasia 
(eastern), Dagestan. Endemic. Described from Dagestan. Type in 
Leningrad. 

Note. T. floccosus Waldst. and Kit. was mistakenly reported for the 
flora of the USSR. It is an endemic of sands along the Danube; it 
differs from T. daghestanicus by the narrowly linear, grooved basal 
leaves, smaller, numerous capitula, and chiefly marginal achenes, which 
are usually smooth in the lower part and tuberculate above, and by the 



175 

presence of a short beak. T. floccosus Waldst. and Kit. belongs to the 
series Dasyrhynchiformes Artemcz. of psammophilous species. 

55. T. borystenicus Artemcz. in Tr. Inst. Bot. Khar'k. Univ. 
II (1937) 47; Klok. in Vizn. Rosl. URSR 590.— 7. brevirostris 
subsp. borystenicus (Artemcz.) C. Rgl. in Scr. Hort. Bot. Univ. Vyt. 
Magni V (1937) 41. — Ic: Artemch. op. cit. Fig. 5; C. Rgl. op. cit. Abb. 
5, 15. 

Biennial. Root robust, cylindrical, hard; root collar densely covered 
with fibers. Plants 30-110 cm high, more or less pubescent; bases, mainly 
of basal leaves and stems below capitual more or less flocculose- 
tomentose, less often almost glabrous. Stems branched, with virgate 
divergent branches. Basal leaves narrower than cauline ones, 2-6 mm 
wide, narrowly linear; upper cauline leaves very short, 3-10 nmn wide 
at base, ovate-cordate, abruptly acuminate, with short recurved cusp; 
cauline leaves pale lilac below, not involute, expanded at base, lamina 
folded or flat, gradually narrowed. Peduncles not thickened below 
175 capitula. Involucral bracts (seven) eight, at fruiting 15-28 mm long, 5- 
7 mm wide, shorter than florets, sometimes violet, as also branches. 
Florets yellow; achenes with pappus 16-19 mm long, about 1 cm long 
without pappus, sulcate along ribs, weakly squamose, sharply scabrous, 
less often almost smooth, gradually narrowed toward both ends, without 
beak; pappus 7-10 mm long, white or greenish-white, as long as 
achene, less often shorter, with annulus of soft hairs at base. Flowering 
May to September. 

Sands of river valleys, steppe regions, and pine forests. — European 
Part: Black Sea Region (lower reaches of the Dnieper and Southern 
Bug, Dzharylagach Island). Endemic. Described from former Kherson 
District. Type in Kiev. 

Note. Plants with weakly scabrous achenes have been found on the 
Aleshki sands; plants up to 110 cm high have been reported from the 
vicinity of Nikolaev and on Dzharylagach Island. C. Regel considers it 
a species of hybrid origin. 

It is possible that after further study T. borystenicus may be 
transferred to another series and, maybe, even to another section {Bessera 
Boriss. or Nikitinia Boriss.). 

Section 11. Sosnovskya Boriss. sect. nov. in Addenda XXVIII, 
724. — Peduncles not thickened below capitula; medium to rather large, 
3.5-5 cm long, on straight peduncles; involucral bracts eight, much 
shorter than florets and as long as outer achenes; florets large, yellow, 
orange-yellow, or pale yellow; achenes large, straight, whitish, gradually 
narrowed into somewhat thick, sometimes indistinct beak shorter than 
body of achene; pappus almost as long as achene with beak. Perennial, 
less often biennial plants, with thick cylindrical roots. 



176 

Type of section: T. reticulatus Boiss. and Huet. 

Series 1. Marginati Kuthath. Kavk. Predst. Tragop. (1957) 26, p. 
p. — Florets yellow; achenes gradually narrowed into somewhat thick, 
indistinct beak shorter than body of achene; pappus dirty-golden, longer 
than achene with beak; root cylindrical, thick; perennial. 

56. T. marginatus Boiss. and Buhse in Nouv. Mem. Soc. Nat. Mosc. 
XII (1860) 136; Kuthath. Kavk. Predst. Tragop. (1957) 26; id. in Fl. 
Azerb. VIII, 505.-7. collinum auct. non DC: Boiss. Fl. or. Ill (1875) 
746, p. p. — T. acanthocarpum Boiss. op. cit. 751 p. p. — A. flexuosus 
Sosn. ex Grossh. Fl. Kavk. IV (1934) 232; Kuthath. in Fl. Gruzii VIII, 
627.— Ic: Kuthath. op. cit. (1957) Plate III, Fig. 10, Map 4. 

Perennial. Root cylindrical, thick; root collar covered with sheaths 
of previous year's leaves. Plants glabrous or glaucous, weakly flocculose, 
10-20(25) cm high. Stems whitish, branched from base or simple. Basal 
176 leaves lanceolate or linear, acuminate, expanded at base; cauline leaves 
much shorter than basal, semiamplexicaul, apiculate, with white border, 
glaucous, often undulate. Peduncles not thickened below capitula. 
Involucral bracts eight, lanceolate, acuminate, flocculose at base, shorter 
than ligulate florets and slightly shorter than achenes with pappus, or as 
long. Capitula ?)-A cm long at fruiting, 2-2.5 cm long at flowering; 
florets yellow. Achenes with pappus about (3)3.7-4 cm long, without 
pappus about 1.5-1.9 cm long, about 2 mm wide, sulcate, whitish, 10- 
ribbed, scabrous, covered with fine subobtuse scales, gradually narrowed 
into thick, sulcate, indistinct beak, half as long as scabrous body of 
achene; pappus about 2.5 cm long, dirty golden, whitish-grayish, slightly 
shorter than achenes with beak. Flowering April to June; fruiting June 
to August. 

Dry slopes, on stony, rubbly, sandy-clayey, clayey and gypsic 
habitats, in the lower and middle mountain zones. — Caucasus: Eastern 
and Southern Transcaucasia, Talysh. Endemic(?). Distribution in 
Iran possible. Described from vicinity of Nakhichevan. Type in 
Leningrad. 

Series 2. Reticulati Boriss. — Flowers yellow, orange-yellow, or pale 
yellow; achenes gradually narrowed into beak, 5-8(10) mm long; 
perennial, rarely biennial plants; capitula medium. 

57. T. reticulatus Boiss. and Huet in Boiss. Diagn. pi. or. ser. II, 3 
(1856) 90; Boiss. Fl. or. Ill, 752; Grossh. Fl. Kavk. IV, 233; Kuthath. 
in Fl. Gruzii VIII, 632; Kavk. Predst. Tragop. (1957) 21; in Fl. Azerb. 
VIII, 504. — T. reticulatus var. stenophyllus Boiss. Fl. or. Ill (1875) 752. — 
T. stenophyllus Boiss. and Huet, op. cit. 89. — Ic: Kuthath. op. cit. (1957) 
Fig. 7. 



177 

Perennial. Root cylindrical, vertical; root collar densely covered with 
reticulate fibers. Stems several, 20-65 cm high, erect, initially tomentose, 
glabrescent, simple, densely leafy. Leaves numerous; basal leaves linear, 
acuminate, many- veined, slightly shorter than stem; cauline leaves 5-12 
mm long at base, amplexicaul, oblong, acuminate, often undulate-crisped, 
not exceeding peduncles, often flocculose along margin. Peduncles not 
thickened below capitula; involucre of eight lanceolate bracts, weakly 
flocculose at base. Capitula 3-4 cm long at flowering; florets bright 
yellow, usually pale lilac on drying, much longer than involucre. Achenes 
with pappus about 3.6 cm long, without pappus about 1.5-1.9 cm long, 
slightly curved, sulcate, finely tuberculate-squamose, scabrous, beak flat, 
0.25-0.5 cm long, somewhat thick, sulcate; weakly swollen at apex and 
with undulate annulus; pappus 1.7-2 cm long, white or grayish. Flowering 
May to June; fruiting June to July. 

Alpine and subalpine meadows, at 1,900-3,000 m. — Caucasus: 
Dagestan, Eastern Western Transcaucasia (rarely), and Southern 
177 Transcaucasia, Talysh. General distribution: Armenia and Kurdistan, Asia 
Minor. Described from vicinity of Erzurum. Type in Geneva. 

Note. Sommier and Levier (1900) reported var. macroglossus Somm. 
and Lev., from Svanetia, which differs from the typical form by the 
larger capitula, florets, and leaves. 

58. T. charadzeae Kuthath. in Zam. po Sist. i Geogr. Rast. Tbil. 
17 (1953) 30; in Fl. Gruzii VIII, 632; Kavk. Predst. Tragop. (1957) 
20.— Ic: Kuthath. op. cit. (1953) Fig. 2; op. cit. (1957), Plate II, Fig. 5. 
Map 1. 

Perennial. Root vertical, cylindrical; root collar densely covered with 
fibers and remnants of old leaves. Plants light green, glabrous or weakly 
flocculose at stem base. Stems 40-60 cm high, solitary or two, erect, 
simple, cylindrical, bearing single capitulum at apices of stems. Basal 
leaves numerous, narrowly linear, long, not exceeding peduncles; cauline 
leaves lanceolate, expanded at base, amplexicaul, long-acuminate. 
Peduncles not thickened below capitula, covered with yellowish 
flocculose pubescence; florets pale yellow. Involucral bracts eight, 
lanceolate, 2.5-2.7 cm long, at fruiting deflexed, often involute, shorter 
than achenes with pappus. Peripheral achenes about 4.2 cm long with 
pappus, straight, without pappus 2.3 cm long, body of achene about 1.8 
cm long, indistinctly ribbed above, indistinctly squamose along ribs, 
gradually narrowed into straight, about 0.5 cm-long beak; pappus about 
1.8 cm long, shorter than achene, yellowish-golden, with undulate annulus 
at base; inner achenes about 3.8 cm long with smooth body, weakly 
sulcate above, but without scales, gradually narrowed into about 0.5 
cm-long beak. Flowering June to July; fruiting July. 



178 

Subalpine meadows. — Caucasus: Eastern Transcaucasia (Georgia, 
Akhalkhevi District). Endemic. Described from Georgia. Type in Tbilisi; 
isotype in Leningrad. 

Note. From T. filifolius Rehm., with which it is united by 
Kuthatheladze (op. cit.) into one series, it is distinguished by wide basal 
leaves and broadly lanceolate cauline leaves (and not filiform basal leaves), 
deflexed, and not erect, involucral bracts, longer pale yellow (and not 
orange-yellow) florets, longer and entirely smooth inner achenes with a 
short beak, and a 14-18 mm-long (not about 1 cm-long) pappus. 

59. T. maturatus Boriss. in Fl. Turkm. VII (1960) 297.-7. praecox 
S. Nikit. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, VII, 12 (1937) 
273, non Focke (1906). 

Biennial. Root cylindrical, v^tical. Plants glabrous, branched below, 
178 leafless above. Stems 50-80 cmiiigh, more or less 3 mm thick. Leaves 
remote, more or less undulate, expanded and amplexicaul at base; lower 
leaves Unear, 7-12(14) nmni wide at base, 18-25 cm long, 5-8(10) mm 
wide in middle; middle leaves 10-12 cm long, about 10 mm wide. 
Peduncles 8-30 cm long, much longer than leaves, 2.5^(6) mm thick 
below capitula. Capitula about 3 cm long at flowering, 3-3.5(5) cm long 
at fruiting; involucral bracts 8-10, lanceolate-linear, thin-acuminate, much 
shorter than florets and slightly shorter than achenes; florets exceeding 
involucre, about 2.5 cm long, pale lilac on drying, thin. Peripheral achenes 
with pappus 3-4.3 cm long, without pappus 1.7-2.5 cm; body of achene 
0.8-1.5 cm long, sulcate, sharply squamose, mainly above with remote 
erect scales; weakly scabrous below, basally drawn out and smooth, 
gradually narrowed toward apex into 0.8-1 cm-long, sulcate beak, almost 
without scales, not swollen at apex and with hairy annulus; pappus dirty 
white, with unequal hairs shorter than achene with beak, 1.3-1.8 cm long. 
Flowering May to June; fruiting May to July. 

Mountains, on slopes, at 1.550-1,650 m. — Soviet Central Asia: 
mountainous Turkmenia. General distribution: Iran. Described from Iran. 
Type in Leningrad. 

Section 12. Dubjanskya Boriss. sect. nov. in Addenda XXVIII, 724. — 
Peduncles strongly thickened below capitula; capitula large, 5-6 cm long, 
solitary, on long peduncles; involucral bracts 8-12; florets yellow; achenes 
with short, about 3 nun long, thick beak; pappus as long as achene or 
slightly longer; biennial, robust, psammophilous plants. 

Type of section: T. sabulosus Krasch. and S. Nikit. 

60. T. sabulosus Krasch. and S. Nikit. in Otch. Pochv.-Bot. Otr. 
Kazakhst. E'ksp. Akad. Nauk SSSR, IV, 2 (1930) 294; Nikit. in Izv. 
Glavn. Bot. Sada XXIX, 5-6, 489; Pavlov. Fl. Tsentr. Kazakhst. Ill, 338; 



179 

Kryl. Fl. Zap. Sib. XI, 2976; C. Rgl. in Berichte Schweiz. Bot. Gesellsch. 
65 (1955) Karte 3; Kovalevskaja in Fl. Uzb. VI, 433.— Ic: Krasch. and 
S. Nikit. op. cit. Fig. 2. 

Biennial. Root long, robust, straight, fusiform, unbranched; root collar 
sometimes covered with fibrous remnants of previous year's leaves. Plants 
40-150 cm high, entirely glabrous. Stems up to 1.5 cm thick, erect, weakly 
sulcate, branched above, densely leafy from base. Lower leaves green, 
linear, almost not expanded at base, 18^0 cm long, 5-10 mm wide, 
often folded, withering at fruiting; middle leaves linear, glaucous, 
semiamplexicaul, up to 10-12(25) mm wide at base, narrowed upward, 
acuminate, 10-25 cm long, sometimes folded; upper leaves few, (3)4-10 
179 cm long, 5-15 mm wide, ovate at base, abruptly narrowed, thin-acuminate, 
sometimes upper leaves absent or persisting up to inflorescence. Peduncles 
clavate below capitula. Capitula 2-10, (3.5)4.5-5.5(6.5) cm long and 1.5- 
3 cm, wide at fruiting; involucral bracts 8-12, 3.5-5 cm long, 5-9 mm 
wide, broadly Unear, gradually narrowed, long-acuminate, shorter than 
florets and achenes, deflexed after fruiting; florets yellow, large. Achenes 
with pappus 3.5-5 cm long, without pappus 1.6-2.5 cm; body of achene 
five-angled, 1.2-1.6 cm long, 1-2.5 mm wide, finely sulcate, finely 
scabrous-tuberculate, with short and about 3 mm long, thick, sulcate beak, 
with hairy annulus at apex; pappus 2.3-2.6 cm long, slightly longer than 
achene with beak, whitish or yellowish, with unequal longer hairs smooth 
at apex. Flowering May; fruiting June. (Plate IX, Fig. 2.) 

Hummocky sands with sparse thickets. — Western Siberia: Upper 
Tobol, Irtysh, Altai; Soviet Central Asia: Aralo-Caspian, Balkhash Region, 
Kyzyl-Kum. General distribution: Dzhungaria-Kashgaria, Mongolia 
(western Mongolia). Described from sands in lower reaches of Sary River. 
Type in Leningrad. 

Economic importance. A fodder plant. 

Section 13. Dasypogon Boriss sect. nov. in Addenda XXVIII, 724. — 
Peduncles not thickened; capitula 3-3.5(4) cm long, numerous; involucral 
bracts five, less often six, slightly shorter than florets and achenes, 
pubescent; florets pale yellow, pale violet on drying; achenes less 
numerous, about five, without beak, somewhat thick, sulcate and 
tuberculate; pappus slightly exceeding achene; perennial or biennial plants, 
spreading and branched from base. 

Type of section: T. kopetdaghensis Boriss. 

61. T. kopetdaghensis Boriss. in Fl. Turkm. VII (1960) 296, 380. 

Perennial or biennial. Root virgate, thickened; root collar sometimes 
branched, covered with brownish remnants of old dead leaves. Plants 
20-25 cm high, spreading, branched from very base. Stems few, erect or 



180 

arcuate, pubescent mainly near peduncles, almost glabrous at flowering, 
finely sulcate; all stems terminating into fertile capitula. Cauline leaves 
narrow, linear, acuminate with filiform apex, semiamplexicaul, expanded 
at base up to 6-8 mm, flocculose-lanate, tomentose mainly near leaf base; 
upper leaves up to 4 cm long, lower up to 20 cm, sometimes folded, 
longer than peduncles. Peduncles not thickened, tomentose near base 
of capitulum. Capitula medium, at fruiting 3-3.5(4) cm long, about 
1-1.5 cm wide in middle; involucral bracts five, less often six, 2.5-3.5 cm 
long (with pappus), lanceolate, broader, at base up to 4-5 mm, narrowed 
toward apex, tomentose at flowering, with sparse pubescence or almost 
180 glabrous at fruiting; florets pale yellow, pale lilac on drying, with five 
violet teeth and dark veins, slightly longer than involucre, about 1 cm 
long, about 1.5 mm wide; with white intertwined hairs at corolla base; 
stamens about 5 mm long, dark violet. Achenes few, about five, somewhat 
thick, without beak; outer achenes with pappus about 3.5 cm long, slightly 
curved, four- or five-ribbed, finely sulcate and sharply tuberculate 
throughout, more sharply sulcate and weakly tuberculate at base, about 
1.5 cm long, narrowed at apex and with hairy annulus; pappus shghtly 
longer than achene, about 2 cm long, with plumose, almost equal hairs, 
with few prominent filiform brownish hairs. Flowering May. (Plate X, 
Fig. 2.) 

Gravelly slopes in wormwood semidesert zone. — Soviet Central Asia: 
mountainous Turkmenia (central Kopetdag, southeastern spurs of Kelyat 
Range). Possibly occurring in Iran. Described from Kopetdag. Type in 
Leningrad. 

Note. The only species of goat's-beard in Turkmenia with a beakless 
achene. From all other species having achenes with or without a short 
beak, it is distinguished by a pappus larger than the achene, involucral 
bracts five, less often six (but not eight), shorter ligulate florets, a small 
number of achenes in the capitulum, and by the sculpture of the achenes. 

Section 14. Nikitinia Boriss. sect. nov. in Addenda XXVIII, 725. — 
Ser. Dasyrhynchiformes Artemcz. in Uch. Zap. Chernov. Univ. I (1948) 
89, p. p. — Peduncles not thickened; capitula medium, 20-45 mm long, 
numerous, on short peduncles, in compound panicles; involucral bracts 
(7)8-12; florets yellow; achenes somewhat thick with or without short 
thick beak; pappus slightly shorter, slightly longer or as long as achene; 
biennial plants with sulcate branched stems, confined to sands and sandy 
soils. 

Type of section: T. dubjanskyi Krasch. and S. Nikit. 

62. T. dubjanskyi Krasch. and S. Nikit. in Otch. Pochv.-Bot. Otr. 
Kazakhst. E'ksp. Akad. Nauk SSSR IV, 2 (1930) 289; S. Nikit. in Izv. 



181 

Glavn. Bot. Sada XXIX, 5-6, 489; Pavlov, Fl Tsentr. Kazakhst. Ill, 338; 
Kryl. Fl. Zap. Sib. XI, 2978; C. Rgl. in Berichte Schweiz. Bot. Gesellsch. 
65 (1955) Karte 2.— Ic: Krasch. and S. Nikit. op. cit. Fig. 1. 

Biennial. Root fusiform, straight, robust, long, unbranched; root collar 
not covered with fibers. Plants 40-150 cm high, wholly glabrous. Stem 
robust, up to 1.5 cm thick at base, straight, smooth or weakly sulcate, 
strongly branched from middle, densely leafy from base. Lower leaves 
green, glabrous, linear, slightly expanded at base, 15-40(50) cm long, 5- 
15 mm wide, folded, later completely drying out; middle leaves glabrous, 
182 glaucous, semiamplexicaul, 8-10-veined, 10-30 cm long, 12-30 mm wide 
at base, gradually narrowed upward to 3-8 mm, often folded, undulate; 
upper leaves much shorter, 4-10 cm long, 5-30 mm wide at base, ovately 
expanded up to 25-30 mm and obtuse, amplexicaul, abruptly narrowed 
at apex, or not narrowed at all. Peduncles not thickened below capitula; 
capitula less numerous (10-70), in compound panicles, 20-45 mm long 
at fruiting. Involucral bracts (7)8-12, broadly lanceolate, 20-30 mm long, 
3-6 wide, narrowed toward apex, glabrous, glaucous, deflexed after 
fruiting and seed shedding; florets light yellow. Peripheral achenes large, 
oblong, with pappus 2.5-3.5 cm long, without pappus 1.4-2 cm, 1.5-2 
mm wide; indistinctly five-angled, almost globose, spinulose-tuberculate, 
ribbed and sulcate, less often smooth, weakly curved, without beak; pappus 
whitish or dirty yellowish, about 1.5 cm long, slightly shorter than achene, 
without hairy annulus at base, one-rowed plumose, with unequal hairs; 
longer hairs not plumose at apex; inner achenes more or less glabrous or 
weakly sulcate. Flowering May to June; fruiting May to August. 

Hummocky, sparsely vegetated sands of western Kazakhstan, less 
often in sandy steppes. — Soviet Central Asia: Aralo-Caspian Region. 
Endemic. Described from sands of Kok-Dzhid on Emba River. Type in 
Leningrad. 

Economic Importance. A food and fodder plant. 

63. T. scoparius S. Nikit. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk 
SSSR, VII, 12 (1937) 262; Artemcz. in Tr. Inst. Bot. Khar'k. Univ. II, 
43, in obs. 

Biennial: Root robust, vertical, straight; root collar covered with 
remnants of old leaves. Plants large, up to 1.5 m high. Stems thick, sulcate, 
strongly branched above, flocculose-tomentose above and at leaf bases. 
Lower leaves linear, weakly expanded at base; middle cauline leaves flat, 
broadly linear above, oblong-lanceolate below, semiamplexicaul, 10- 
15(30) cm wide, abruptly narrowed toward apex, more or less undulate, 
sometimes involute; upper leaves wide, reduced, oblong or lanceolate- 
oblong in lower part, abruptly narrowed above, semiamplexicaul. 
Peduncles not thickened below capitula, 1-2(3) mm thick. Capitula 



182 




181 Plate X. 

1 — Tragopogon wolgensis S. Nikit.; 2 — T. kopetdaghensis Boriss. 



183 

numerous on short peduncles in paniculate inflorescence, 1.5-2(2.5) cm 
long at flowering, 2.5-3.5 cm long at fruiting. Involucral bracts 8-10, 
lanceolate, expanded at base, narrowed toward apex, distinctly shorter 
than florets and achenes— by (2)4-8 mm; florets light yellow. Peripheral 
achenes including pappus about 2.4 cm long, excluding pappus about 1 
cm, brownish, curved, sulcate, indistinctly five-angled, finely squamose 
183 scabrous, gradually narrowed into thick and short, indistinct, sulcate, 
squamose, 1-3 mm-long beak; pappus slightly longer than achene, or 
almost as long. Flowering June to July; fruiting July to August. 

Steppe meadows, mountain slopes, sometimes forming thickets on 
wastelands. — Soviet Central Asia: Dzhungaria-Tarbagatai, Balkhash 
Region. Endemic. Described from vicinity of Lepsinsk. Type in Leningrad. 

Note. Similar to T. dubjanskyi, Krasch. and S. Nikit., from which it 
is distinguished by the linear leaves and smaller capitula, as well as by 
the form and sculpture of the achenes. 

Economic Importance. As a fodder plant relished by cattle. 

64. T. ruthenicus Bess, ex Krasch. and S. Nikit. in Otch. Pochv.- 
Bot. Otr. Kazakhst. E'ksp. Akad. Nauk SSSR IV, 2 (1930) 292; in Izv. 
Glavn. Bot. Sada, XXIX, 487; Fl. Yugo-Vost. VI, 443; Pavlov, Fl. Tsentr. 
Kazakhst. HI, 337; C. Rgl. in Scr. Hort. Bot. Univ. Vyt. Magni V, 43 
and in Berechte Schweiz. Bot. Gesellsch. 65 (1955) Karte 1. — T. villosum 
auct. non L.: Pall. Reise V (1778) 332, p. p.— r. canus auct. non Willd.: 
Bess. Enum. Fl. Volh. (1822) 30. — T. ruthenicus subsp. typicus C. Rgl. 
op. cit. (1937) 43, 44; and op cit. (1955) Karte 1. — Pteropogon ruthenicus 
CAM. ex Claus in Goebel, Reise in d. Stepp. Sudl. Russl. 2 T.(1838) 
284, nomen nudum.— Ic: S. Nikit. in Fl. Yugo-Vost. VI, 443, Fig. 715; 
C. Rgl. op. cit. (1937) Abb. 7, 18. 

Biennial. Root robust, long, vertical; root collar densely covered with 
fibrous remnants of old leaves. Plants 30-80(115) cm high, flocculose- 
tomentose, especially above, on young branches and in leaf axils. Stems 
erect or slightly flexuous, usually robust, thick, highly sulcate, usually 
branched above, densely leafy from base, fistular below. Lower basal 
leaves usually expanded at base, 20^0 cm long, 4-12 mm wide, often 
folded, withering at seed maturation, acute; middle leaves semiamplexicaul, 
9-15(18) cm long, expanded at base, 8-25 mm wide, gradually narrowed 
upward and glabrous in this part, conduphcate along keel, sometimes 
undulate; uppermost leaves reduced, 2-6 cm long, 6-10 mm wide at base, 
spatulate, narrowed above, with long, arcuate cusp, usually flocculose- 
tomentose. Peduncles not thickened below capitula, 1-2(3) mm thick. 
Capitula 3-30, at fruiting 20^0 mm long, 10-15 wide. Involucral bracts 
7-12, at fruiting (18)25-35 mm long, 4-5 mm wide, more or less 
pubescent, especially very young ones, acuminate, glaucous from 



184 

flocculose tomentum, slightly shorter as long as florets but shorter than 
achenes; florets light yellow. Peripheral achenes 1.5-3 cm long with 
pappus, without pappus 1.2-1.5(2) cm, somewhat thick, about 1-1.5 mm 
wide, shallow-sulcate, with 8-12 alternating smooth and finely tuberculate 
ribs, straight or somewhat curved, without beak, sometimes smooth in 
184 upper part, not spinulose; pappus 1.5-1.8 cm long, as long as or longer 
than achene, sometimes reddish with hairy annulus at base. Flowering 
May to August (September). 

Hummocky sands and slopes and in hollows, less often at the top of 
hummocks, dry sandy substrates, in river valleys. — European Part: Lower 
Volga, Lower Don; — Soviet Central Asia: Aralo-Caspian Region. Endemic. 
Described from Caspian steppes. Type in Leningrad. 

Note. The species is close to T. dubjanksyi Krasch. and S. Nikit., 
from which it differs by having less numerous (3-30, not 10-70) capitula, 
a smaller size, less branching, smaller involucres and achenes, leaves 
narrower at the base and flocculose-tomentum on the stem, leaves, and 
involucres. 

Economic Importance. A fodder plant. 

65. T. donetzicus Artemcz. in Tr. Inst. Bot. Khar'k. Univ. II (1937) 
49; Klok. in Vizn. Rosl. URSR, 590; C. Rgl. in Berichte Schweiz. Bot. 
Gesellsch. 65 (1955) Karte 2. — Ic: Artemcz. op. cit. Fig. 6. 

Biennial. Plants 10-40 cm high, more or less densely pubescent, 
densely flocculose below capitula and on young branches. Stems solitary, 
or sometimes several, often pyramidally branched from base. Upper leaves 
20-35 mm long, not shorter than 15 mm, similar to middle cauUne leaves, 
not involute; basal leaves expanded at base but narrower than cauline 
leaves, narrowly linear, about 3 mm wide, narrowed upward and folded; 
cauline leaves reduced, expanded at base, 6-7 mm wide, folded, sometimes 
flat, acute, deflexed and bent. Involucral bracts seven or eight, 15-20 
mm long at fruiting, 5-10 mm wide, shorter than florets, with wide white 
stripe along margin, densely or sparsely pubescent, alternating with each 
other. Peduncles not thickened below capitula, 5-10 mm thick. Capitula 
at fruiting 10-19 mm wide; florets yellow. Achenes 10-14 mm long, 
gradually narrowed toward apex, weakly scabrous; peripheral achenes 
sparsely pubescent, without or almost without beak; pappus reddish-gray, 
10-11 mm long, with glabrous, annulus at base. Flowering June to 
September. 

Sands in river valleys, dry sands, and less often in steppe areas. — 
European Part: ? Black Sea Region, Lower Don (middle reaches of 
Donets). Endemic. Described from Kharkov Region near village of 
llichevka. Type in Kiev. 

Note. It differs from T. tanaiticus Artemcz. by having a shorter stem. 



185 

narrower leaves and a glabrous, thickened annulus at the base of the 
pappus. From T. ucrainicus Artemcz. it differs by having beakless achenes. 

Section 15. Chromopappus Boriss. sect, nova in Addenda XXVIII, 
725. — Peduncles clavately thickened below capitula, short; capitula large; 
involucral bracts six to eight, one and one-half to two times as long as 
185 florets; florets purple, pinkish, or violet; achenes with somewhat thick, 
straight, almost smooth beak, longer or slightly shorter than gradually or 
abruptly expanded body of achene; pappus colored, dirty violet, almost 
as long as achene. Biennial plants. 

Type of section: T. coloratus C.A.M. 

66. T. coloratus C.A.M. Verzeichn. Pfl. Cauc.(1831) 61; DC. Prodr. 
VII, 12; Ldb. Fl. Ross. II, 2, 785; Boiss. Fl. or. Ill, 746; Grossh. Fl. 
Kavk. IV, 230; Kuthath. in Fl. Gruzii VIII, 620; Kavk. Predst. Tragop. 
(1957) 41; in Fl. Azerb. VIII, 508; Kitam. Fl. Afghan. (1960) 452.-7. 
ruber auct. non S.G. Gmel.: Grossh. Fl. Kavk. (1934) 230.— Ic: Kuthath. 
op. cit. (1957) Plate V, Fig. 10, Map 7. 

Biennial. Root vertical, thick, sometimes branched; root collar covered 
with remnants of old leaves. Plants 7-30(60) cm high. Stems glabrous, 
densely leafy, branched or simple, erect. Basal leaves long, narrow; cauline 
leaves semiamplexicaul, expanded at base, flat, lanceolate-linear or 
narrowly linear, long-acuminate, undulate, often exceeding peduncles. 
Peduncles, especially at fruiting, strongly clavately thickened, short, often 
drawn out below capitula. Capitula solitary on each peduncle, 4-4.5 cm 
long on fruiting. Involucral bracts six to eight lanceolate-linear, narrowed 
towards apex, acuminate, expanded at base, and membranous along 
margin, especially at fruiting; florets purple, pinkish, or violet, half to 
two-thirds as long as involucre. Achenes including pappus 5.3 cm, 
excluding pappus 2.7 cm long; body of achene 1 .5 cm long, finely sulcate, 
with 10 ribs, scabrous from small dense scales or tubercles; gradually 
narrowed into somewhat thick, 1.2 cm-long, straight, glabrous, almost 
smooth beak, without hairy annulus at apex and clavately thickened; 
pappus 2.4 cm long, dirty violet, with few longer hairs, almost as long as 
achene with beak. Flowering May to June; fruiting June to July. 

Dry grassy and stony slopes in lower and middle mountain zones. — 
Caucasus: Eastern and Southern Transcaucasia, Talysh. General 
distribution: Balkans-Asia Minor, Iran. Described from Zuvant. Type in 
Leningrad. 

Note. In the Nakhichevan ASSR, var. floccosus Sosn. (Kuthath. 
[1957] Kavk. Predst. Tragop., 42), with flocculose stem and leaves, is 
recognized. 

67. T. nachitschevanicus Kuthath. in Zam. po Sist. i Geogr. Rast. 



186 

Tbil. 15 (1949) 96; Kavk. Predst. Tragop. (1957) 59; in Fl. Azerb. VIII, 
511.— Ic: Kuthath. op. cit. (1957) Plate IX, Fig. 30, Map 10. 

Biennial. Root vertical; root collar often covered with remnants of 
previous year's leaves. Plants glabrous, 40(60) cm high. Stem erect, 
branched, densely leafy, finely sulcate. Lower leaves expanded at base, 
semiamplexicaul, abruptly narrowed, long-acuminate; cauline leaves up 
186 to 2 cm wide at base; all leaves narrowed toward apex, long-acuminate, 
semiamplexicaul. Capitula up to 7 cm long, solitary. Peduncles strongly 
thickened upward; involucral bracts six to eight, expanded at base, long- 
acuminate, longer than achenes including pappus, at base glabrous; florets 
purple. Peripheral achenes large, somewhat curved, about 6 cm long 
with pappus, 3.2 cm long without pappus; body of achene about 1.3 cm 
long, 3 mm wide, with three grooves on outer side and two on inner, 
with five prominent, sometimes winged, squamose-tuberculate ribs, at apex 
abruptly narrowed into smooth, white, somewhat thick, five-angular, 1 .9 
cm-long beak, one and one-half times as long as body of achene, 
somewhat thickened at apex, not clavate, with hairy annulus; pappus 2.7 
cm long, purple above, hairs unequal. Inner achenes larger, quite different 
from outer, straight, shallow-sulcate, finely transversely squamose; body 
of achene gradually narrowed into long beak. Flowering May to June; 
fruiting May to July. 

Screes and gravelly slopes in the middle mountain zone. — Caucasus: 
Southern Transcaucasia, Talysh. Endemic. Described from Nakhichevan 
ASSR, village of Saltakh. Type in Leningrad; isotype in Tbilisi. 

Section 16. Rubriflori Boiss. Fl. or. Ill (1875) 745, p. p. — set. Collini 
Kuthath. Kavk. Predst. Tragop. (1957) 25, p. p. — Peduncles below capitula 
not thickened, weakly or somewhat strongly thickened; capitula solitary, 
medium to large, on long straight peduncles; involucral bracts 8-10, shorter 
or almost as long as ligulate florets; florets purple, pink, or violet; achenes 
thicker below, tuberculate-squamose, sulcate, sometimes almost winged, 
gradually narrowed into somewhat thick, straight beak; pappus as long as 
or slightly shorter or longer than achenes; plants perennial, weakly 
branched, 5^0 cm high, usually with numerous leaves crowded in basal 
part; roots cylindrical, thick. 

Type of section: T. ruber S.G. Gmel. 

Series 1. Collini (Kuthath.) Boriss. comb. nov. — sect. Collini Kuthath. 
Kavk. Predst. Tragop. (1957) 25, p. p. Short, 5-25 cm high, plants. 
Peduncles not thickened; involucral bracts eight, usually shorter or almost 
as long as florets; capitula large, 4.5-4.8 cm long at fruiting, solitary; 
florets pink, reddish, or purple; achenes with thick, almost smooth beak, 
gradually broadened into sharply scabrous body of achene; pappus 
sometimes dirty violet, shorter than achene. 



187 

68. T. coUinus DC. Prodr. VII (1838) 115; Ldb. Fl. Ross. II, 2, 789; 
Boiss. Fl. or. Ill, 746 (excl. syn. T. marginatum Boiss. and Buhse); Grossh. 
Fl. Kavk. IV, 230; Rech. f. Symb. Afghan. II, 189; Kuthath. Kavk. Predst. 
Tragop. (1957) 28; in Fl. Azerb. VIII (1961) 505; Kitam. Fl. Afghan. 
187 (1960) 452. — T. roseus auct. fl. cauc. noh Trevir.: C.A.M. Verzeichn. 
Pfl. Cauc. (1831) 61. — T. mutabile auct. fl. cauc. non Trevir.: Hohen. in 
Bull. Soc. Nat. Mosc. XI(1838) 273.— Ic: Rech. op. cit. fig. 128; Kuthath. 
op. cit. (1957), Plate III, Fig. 11, Map 4. 

Perennial or biennial. Root cylindrical, vertical, thick; root collar 
covered v^'ith fibers and remnants of old leaves. Plants flocculose, glaucous. 
Stems, 5-25 cm tall, solitary, simple, less often weakly branched, usually 
each with solitary capitulum, densely leafy. Basal leaves numerous, 
expanded at base, semiamplexicaul, lanceolate-linear or filiform, flat or 
undulate and involute, gradually narrowed upward, with white border, 
almost as long as stem or sometimes slightly longer; cauhne leaves whitish 
toward base and expanded into sheath, lanceolate-linear, undulate, usually 
curved. Peduncles not thickened below capitula. Involucral bracts eight, 
shorter or almost as long as florets, narrowly lanceolate; capitula 2.5-3 
cm long at flowering, 4.5^.8 cm long at fruiting; florets pink, reddish, 
or purple, usually longer than involucre, Achenes with pappus 4.8 cm 
long, 2.5 cm without; body of achene excluding beak 1 .5 cm long, about 
2 mm wide, scabrous and with sharp ribs in middle, smooth below, 
gradually narrowed above into thick, about 1 cm-long, smooth beak two- 
thirds as long as body of achene, without apical thickening and hairy 
annulus; pappus 1.8-2 cm long, usually dirty violet. Flowering April to 
May; fruiting May to June. 

Dry slopes of hills, dry stony slopes, and coastal sands. — Caucasus'. 
Eastern and Southern Transcaucasia. General distribution: Iran, 
northwestern Afghanistan. Described from vicinity of Baku. Lectotype in 
Leningrad. 

Note. Kuthatheladze (op. cit, recognizes var. coeruleus, Kuthath, with 
Ught blue Ugulate florets, which are much longer than the involucral bracts; 
plant collected from Zuvant. T. collinus DC. differs from T. ruber Gmel., 
described from the vicinity of Astrakhan, by having involucral bracts 
that are half as long as the florets. It also differs from T. mutabile Jacq. 
by having a more dense pubescence, undulate leaves, and smaller size. 
The report of T. collinus DC. for Alai probably relates to T. alaicus S. 
Nikit. 

Series 2. Rubri Boriss. — Peduncles not thickened or weakly thickened 
at fruiting; capitula solitary at apices of stem and branch; 4-6 cm long; 
involucral bracts 8-10, usually shorter than florets; achenes with somewhat 
thick, sulcate beak, gradually or abruptly expanded into thicker sharply 



188 

scabrous and deeply sulcate, sometimes winged, body of achene, longer 
than beak; pappus yellowish or dirty violet, as long or slightly longer 
than achene. 

69. T. ruber S.G. Gmel. Reise II (1774) 198; DC. Prodr. VII, 114; 
Boiss. Fl. or. 746; Ldb. Fl. Ross. II, 2, 789; S. Nikit. Fl. Yugo-Vost. VI, 
445; Schmalh. Fl. II, 145; Pavlov, Fl. Tsentr. Kazakhst. Ill, 336; C. Rgl. 
in Berichte Schweiz. Bot. Gesellsch. 65 (1955) Karte 2. — T. mutabile 
auct. fl. ross. non Jacq.: Schmalh. op. cit. 145. — T. roseus Trevir. in 
Mag. Ges. Natutf. Fr. Berl. VII (1816) 152.— Ic: Ldb. Ic. fl. ross. I, t. 
30. 

Perennial. Root vertical, cylindrical, robust; root collar covered with 
old leaves. Plants 8-25(40) cm high, glaucous, young plants arachnoid, 
later glabrous and dark glaucous. Stems straight, erect, branched below 
or simple, flocculose-tomentose in leaf axils and below capitula. Basal 
and lower leaves linear, 10-20 cm long, (5)7-15(20) mm wide, expanded 
at base; cauline leaves lanceolate, all entire; middle leaves shorter and 
broader than lower, lanceolate or linear-lanceolate, with broad 
semiamplexicaul base, more or less undulate, less often almost flat, 
acuminate and sometimes involute, scarious along margin. Peduncles long, 
at fruiting not or weakly thickened, 2-4(7) mm in thick. Capitula solitary 
at apices of stem and branch, large, 4.5-6 cm long at fruiting, 3-5 mm 
long at flowering; involucre of 8-10 bracts, shorter or almost as long as 
florets and achenes. Involucral bracts 8-12 mm wide, 25-30 mm long, 
arachnoid-flocculose, glabrescent, smooth, lanceolate, long-acuminate. 
Florets pink-violet, purple, or light violet; peripheral florets one and one- 
half times as long as involucre. Peripheral achenes with pappus 4-5 cm 
long, exceeding involucre, without pappus 2.1-2.15 cm long; body of 
achene 1.4-1.6 cm long, about 1.5 mm wide, sulcate, densely covered 
with white, obtuse scales, sometimes weakly sculptured, gradually 
narrowed into somewhat thick, sulcate beak weakly squamose below, 
shorter than achene, 0.7-1.0 cm long, without apical swelling; pappus 
dirty violet or yellowish, about 2.5 cm long, almost as long as achene 
including beak, usually without hairy annulus at base. Flowering April to 
June; fruiting May to August (Plate XI, Fig, 2). 

Sandy and clayey steppe areas, gentle and stony slopes in desert- 
steppe zone; now and then on scarps, sands, dry sunny slopes, sometimes 
a weed in gardens — European Part: Lower Volga; Soviet Central Asia: ? 
Aralo-Caspian, Balkhash Region; Western Siberia: Irtysh. General 
distribution: ? Dzhungaria-Kashgaria. Described from vicinity of 
Astrakhan. Type lost. 

70. T. marginifolius Pavl. in ByuU. Mosk. Obshch. Isp. Prir. XL VII, 
2 (4 II 1938) 83; Pavlov in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, 



189 

IX, 2 (1941) 52; Kovalevskaja in Fl. Uzb. VI, 432. — T. gonocarpum S. 
Nikit. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR VII, 12 (22 V 
1938) 266.— r. gonocarpus (S. Nikit.) Stank, in Stank i Tal. Opred. Vyssh. 
Rast. (1949) 688. — T. marginatum Pavl. in Byull. Mosk. Obshch. Isp. 
Prir. XLII, 2 (1933) 135, non Boiss. and Buhse (I860).— r. abolinii ssp. 
gonocarpum S. Nikit. in Fl. Yugo-Vost. VI (1936) 445.— T. abolinii M. 
Pop. and Vved. ex C. Rgl. in Berichte Schw. Bot. Gesellsch. 65 (1995) 
Karte 2 nom. — Ic: Pavlov, op. cit. (1933) Fig. 2; Kovalevskaja, op. cit. 
Plate XLIII, Fig. 4. 
189 Perennial. Tap root vertical, thick. Plants 10-40 cm high, branched 

from base or middle or stems simple, young branches sometimes reddish, 
glaucous, sometimes flocculose-tomentose, glabrescent. Stem rather thick, 
sulcate, glabrous. Basal and cauline leaves broadly lanceolate, 10-20(30) 
mm wide, their maximum width below middle, semiamplexicaul, many- 
veined, often with undulate cartilaginous whitish border, more or less 
undulate, somewhat attenuate toward apex, thin; upper leaves reduced, 
oblong-lanceolate to ovate-oblong. Peduncles below fertile capitula weakly 
thickened or not, 3-5(7) mm thick, fistular. Capitula solitary, large, 20- 
40 mm long at flowering, 40-60 mm at fruiting. Involucral bracts eight, 
lanceolate, with more or less prominent midrib, acuminate, shorter than 
florets and achenes; florets pink- violet. Peripheral achenes with pappus 
about 4 cm long, without pappus about 2 cm, sharply five-angled and 
sulcate, imbricate-squamose, sometimes winged along ribs, yellowish, 
abruptly narrowed into somewhat thick, angular, ribbed, about 6 mm- 
long beak, sharply squamose below and one-third to half as long as achene, 
less often as long, not swollen at apex; pappus about 2.5 cm long, 
yellowish or reddish-brown, somewhat longer or almost as long as achene 
with beak. 2n = 12. Flowering May to June; fruiting June to July. 

Stony peaks and screes on dry south slopes, in semidesert, on butte- 
like elevations. European Part: Lower Volga (Bogdo, Chopchachi); Soviet 
Central Asia: Aralo-Caspian Region (Mugodzhary, Lake Inder, Aktau, 
Emba River, Ust-Urt near Aral Sea), Balkhash Region (Chu-Ili Mountains); 
Tien Shan (Talass Alatau, Karatau Range). Endemic. Described from 
Boroldai Mountains (Karatau Range). Type in Moscow. 

Note. It differs from T. ruber S.G. Gmel. by having acutely angular, 
deeply sulcate, wing-ribbed achenes, cartilaginously bordered leaves, and 
shorter involucral bracts at fruiting. 

71. T. conduplicatus S. Nikit. in Bot. Mat. Gerb. Bot. Inst. Akad. 
Nauk SSSR VI 12 (1937) 265.— T. rubrum auct. fl. turkest. ex parte non 
S.G. Gmel. 

Perennial. Stem slender, short, 10-17 cm long, simple or branched 
below, solitary. Lower leaves narrowly linear, flat, conduplicate; middle 



190 

leaves strongly expanded at base 3-6(12) mm wide, semiamplexicual, 
narrowed into about 10 cm long, linear-lanceolate lamina, usually 
conduplicate; upper leaves undeveloped or reduced. Peduncles not 
thickened below fertile capitula, 2-3 mm thick. Capitula at fruiting 30- 
40 mm wide. Involucral bracts eight, noticeably shorter than florets and 
achenes, less often longer than florets; involucral bracts 2-3 cm long, 
linear-lanceolate, acuminate; florets pink, violet on drying. Penpheral 
achenes about 4.5 cm long, excluding pappus 2.2 cm, five-angled, deeply 
sulcate distinctly sculptured, obtusely squamose throughout, gradually narr- 
190 owed into about 5 mm-long, somewhat thick, sulcate beak, without swollen 
apex and stripes, smooth above; pappus about 25 mm long, as long as 
achene or with beak slightly longer. Flowering May to July; fruiting May 
to August. 

Rare on stony and rubbly mountain slopes. — Soviet Central Asia: 
Pamiro-Alai, mountainous Turkmenia. Endemic. Described from Zeravshan 
Range. Type in Leningrad. 

Note. It differs from T. ruber S.G. Gmel. by having narrow, mostly 
folded leaves, straight and very long, deeply sulcate achenes, and slender 
stems. The involucres in the Kopetdag specimens exceed the florets, and 
the leaves are long, filiform-linear. 

72. T. montanus S. Nikit. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk 
SSSR VII, 12 (1937) 270. —T. rubrum auct. fl. turkest., non S.G. Gmel. 
pro parte. 

Perennial. Stems thick, sulcate, usually branched only below, densely 
leafy, 25-30 cm high. Leaves straight, flat, wide, many-veined; lower 
leaves linear, middle ones linear-lanceolate, long-acuminate, (5)7-15(20) 
mm wide below, semiamplexicaul. Peduncles fistular below fertile capitula, 
not thickened, 3-5 mm thick, sometimes weakly thickened up to 7 mm. 
Capitula 30-40(50) mm long with florets, 4.5-5.5(6) cm long at fruiting. 
Involucre of eight bracts, slightly shorter or almost as long as florets and 
achenes; involucral bracts lanceolate, acuminate, about 4 cm long; florets 
pink-violet. Peripheral achenes about 5 cm long, excluding pappus 2.5 
cm long, without beak 1 .5 cm long, obtusely five-angled, deeply sulcate, 
obtusely squamose-scabrous above, usually indistincdy sculptured below, 
gradually narrowed into about 1 cm-long, thick, sulcate beak, shorter than 
achene, without apical swelling; pappus about 2 cm long, almost as long 
as achene including beak. Flowering May; fruiting May to June. 

Single plants found on dry stony slopes. — Soviet Central Asia: 
Dzhungaria-Tarbagatai, Tien Shan, Pamiro-Alai, mountainous Turkmenia. 
General distribution: Possibly occurring in neighboring parts of Iran. 
Described from Tien Shan. Type in Leningrad. 



191 

Note. S. Nikitin (op. cit. 271) segregates T. montanus ssp. 
tianschanicus S. Nikit., which is characterized by the involucral bracts 
being shorter than the pink-violet florets and the achenes, remote leaves, 
a sometimes shortened stem, the color of the corolla, and a pappus shorter 
than the achene. 

Series 3. Longipapposi Boriss. — Peduncles not thickened; capitula 
large and medium, 4-6 mm long at fruiting; involucral bracts eight, shorter 
than florets and achenes; florets pink-violet; achenes with somewhat 
slender beak gradually widening to body, beak half to two-thirds as long 
as achene; pappus much longer than achene with beak. 

191 73. T. alaicus S. Nikit. in Bot. Mat. Gerb. Bot. Inst Akad. Nauk 

SSSR VII, 12 (1937) 268.-7. rubrum auct. fl. As. Med non S.G. Gmel.— 
T. collinus auct.: B. Fedtsch. Rast. Turk. (1995) 760. 

Perennial. Plants 40-60 cm high; stem sulcate, branched below. 
Leaves narrowly linear to linear-lanceolate; upper leaves reduced, 3-5 
cm long, broader in lower part, with oblong-elliptical base, 5-7(12) mm 
wide, acuminate, more or less finely scarious along margin, sometimes 
slightly undulate. Peduncles with capitula not thickened, 2-3 mm thick. 
Capitula about 4-5 cm long at fruiting. Involucral bracts eight, lanceolate- 
linear, long-acuminate, somewhat shorter than florets and achenes, about 
3 cm long, about 4 mm wide at base; florets pink-violet longer than 
involucral bracts. Peripheral achenes with pappus about 5.8 cm long, 
excluding pappus 2.7 cm, about 1.5-2 cm long without beak, shallow- 
sulcate to ribbed and finely and imbricately squamose, gradually narrowed 
into white, about 0.7-1.2 cm-long, almost smooth beak finely sulcate at 
base, not thickened at apex and with hairy annulus at pappus base; pappus 
yellowish-white, about 3 cm-long, longer than achene including beak. 
Flowering May to June; fruiting June to July. 

Mountain slopes. — Soviet Central Asia: Pamiro-Alai (Alai Range). 
Endemic. Described from Alai Range between Chirchik and Gulcha in 
Kirgizia. Type in Leningrad. 

Note. It differs from T. ruber S.G. Gmel. by the shape of the leaves, 
the involucre being shorter than the florets and the achenes, and the pappus 
being longer than the achenes, as well as by the shape and sculpture of 
the achenes. 

74. T. gaudanicus Boriss. in Fl. Turkm. VII (1960) 298, 381—7. 
rubrum auct. fl. As. Med. non S.G'. Gmel. — Ic: Fl. Turkm. VII, Plate 
XLV, Fig. 2. 

Perennial or biennial. Root virgate, weak, slightly thickened in upper 



192 

part; root collar covered with remnants of old leaves. Plant short, glabrous. 
Stems 10-25 cm high, slender, somewhat ascending, simple or weakly 
branched, densely leafy. Cauline leaves lanceolate or oblong-lanceolate, 
with maximum width in lower third, up to 1 cm wide, semiamplexicaul, 
acuminate. Peduncles exceeding leaves, 3-10 cm long, not' thickened 
below capitula, glabrous, finely sulcate. Capitula solitary at apices of stems 
about 2.5 cm long at flowering, about 5 cm long at fruiting, narrow, 
about 1 cm wide. Involucre of eight lanceolate, acuminate bracts, not 
longer than florets and achenes; florets violet. Achenes about 5 cm long, 
excluding pappus but including beak about 2 cm long; beak about 1 cm 
long, slender, as long as body of achene, finely sulcate; weakly scabrous 
at base; achenes sulcate and scabrous from very small sparse scales; pappus 
about 3 cm long, dirty yellow, with intertwined hairs at base. Flowering 
April to May; fruiting May. 
192 Dry stony mountain slopes. — Soviet Central Asia: mountainous 

Turkmenia. General distribution: may occur in Iran. Described from 
Kopetdag Range. Type in Leningrad. 

Note. It differs from T. montanus S. Nikit. by the better developed 
stem and the leaves, which are not crowded at the base of the stem, the 
form and sculpture of the achenes, and the relationship of the length of 
the achene, pappus, and beak. In the Flora of Turkmenia errors have 
crept in, and the lines are mixed up in the description of T. gaudanicus 
Boriss. 

Series 4. Brevipapposi Boriss. Peduncles not thickened; capitula 
solitary, on erect peduncles, 3.5^ cm long at fruiting, 2-3 cm long with 
florets; involucral bracts eight, shorter than florets and achenes; florets 
pinkish violet; achenes with somewhat thick, almost smooth beak, with 
distinct deeply sulcate and sharply squamose body; pappus reddish-brown, 
densely plumose-hairy, shorter than achene; stems rather tall, slender. 

75. T. elongatus S. Nikit. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk 
SSSR VII, 12 (1937) 269; Kryl. Fl. Zap. Sib. XI, 2974.-7. rubrum auct;. 
mult, non S.G. Gmel. p. p. 

Perennial. Plant 10-35 cm high. Root virgate, vertical, thickened; 
root collar usually branched, covered with remnants of previous year's 
leaves. Stems slender, sulcate, erect, or weakly flexuous, simple or 
branched from base. Leaves erect, linear, oblong; middle cauline leaves 
rhomboidal below 5-8(10) mm wide, narrowed into narrowly Unear lamina 
above, narrowly scarious along margin; upper leaves reduced. Peduncles 
not thickened below capitula, 2-A mm thick. Capitula 2-3 cm long at 
flowering, 3.5^ cm at fruiting. Involucre of eight bracts, shorter than 
florets and achenes; florets pinkish-violet. Peripheral achenes almost 



193 




'93 Plate XI 

' — Tragopogon bjelorussicus Artemcz.; 2 — T. ruber S.G. Gmel. 



194 

straight, 3.5-3.7 cm long including pappus, 2 cm long without pappus, 
about 2 mm wide, about 1.3 cm long excluding beak, obtusely five-angled, 
sulcate, with acute and dense small tubercles or scales, narrowed into 
sulcate, ribbed, somewhat thick, almost smooth, about 0.7 cm-long beak, 
with hairy annulus at apex; pappus brownish, with dense long hairs, 
shorter than achenes including beak, 1.6-1.8 cm long. Flowering May to 
June; fruiting June to July. 

Mountain slopes, in steppe vegetation on sandy-gravelly, clayey and 
stony soils. — Western Siberia: Irtysh; Soviet Central Asia: Balkhash Region 
(Zaisan Kettle hole, Dzhungaria-Tarbagatai, Tien Shan. General 
distribution: Dzhungaria-Kashgaria. Described from vicinity of village of 
Kenderlyk in Zaisan Kettle hole. Type in Leningrad. 

Note. It differs from T. ruber S.G. Gmel. by having linear, straight, 
elongated leaves, that are basally rhomboidal and very narrow above; in- 
194 volucral bracts shorter than the florets and the achenes; a reddish-brown, 
densely plumose pappus that is shorter than the achene, and tall stems. 

Series 5. Asiatici Boriss. Peduncles more or less thickened; plants 
short, densely leafy in basal part; capitula 4-6 nmi long; in volucral bracts 
eight, almost as long as florets but shorter than achenes; achenes with 
somewhat thick smooth beak, almost as long as or shorter than body of 
achene; pappus brownish-red or yellowish, almost as long as achene. 

76. T. malikus S. Nikit. i Bot. Mat. Gerb. Bot. Inst. Akad. Nauk 
SSSR VII, 12 (1938) 267.— Zakir. Fl. i Rast. Zeravsh. 2, 419; 
Kovalevskaja in Fl. Uzb. VI, 431. — T. ruber var. latifolius M. Pop. and 
Vved. ex Lapin, Opr. Rast. Tashkent. Oazisa I (1933) 114.— T. latifolius 
M. Pop. in Tr. Uzb. Gos. Univ. N. S. 27, Biol. 14 (1941) 94; Zakir. op. 
cit. 420. — T. rubrum auct. fl. turk. pro parte, non S.G. Gmel. — Ic: 
Kovalevskaja in Fl. Uzb. Plate XLIII, Fig. 2. 

Perennial. Root cylindrical; root collar covered with remnants of old 
leaves. Plants glabrous, glaucous-green, 20-30 cm high. Stems branched 
or simple, sulcate, with short intemodes, rather densely leafy. Basal leaves 
linear, not longer than inflorescence, hairy along margin, thin acuminate; 
middle cauline leaves narrow, linear-lanceolate, with maximum width in 
lower part, 5-10, very rarely 20 mm wide, slightly narrowed near base, 
more or less sinuate and scarious along margin; upper leaves linear- 
lanceolate to lanceolate and oblong, base up to 10-20 mm, much shorter 
than basal and lower cauline leaves, thin-acuminate. Peduncles strongly 
leafy, somewhat thickened below fertile capitula, 3-4(5) mm thick. 
Capitula large, cylindrical, 40-60 mm long at fruiting, 15-20 mm wide 
(dry). Involucral bracts lanceolate, acuminate, with broad bases, almost 
as long as florets but shorter than achenes with pappus, pubescent at base 



195 

at fruiting, also pubescent at apex at flowering; florets pink-violet. 
Peripheral achenes straight, with pappus about 5 cm long, without pappus 
2.6 cm long, body of achene about 1.5 cm long, five-angled, sulcate, 
finely and sharply squamose, sometimes weakly sulcate, gradually 
narrowed into somewhat thick, smooth or weakly sulcate, about 1.5 cm- 
long beak, not swollen at apex below pappus and with hairy annulus; 
pappus about 2 cm long, its individual hairs not plumose, up to 2.4 cm 
long, reddish-brown with violet tinge, densely plumose-villous. Flowering 
April to May; fruiting May to June. 

Foothill zone. — Soviet Central Asia: Pamiro-Alai (Karatau, Malik, 
Pistalitau mountains). Endemic. Described from Mt. Malik. Type in 
Tashkent. 

Note. It differs from T. marginifolius Pavl. by its narrow leaves, 
cylindrical capitula, and straight achenes. 

195 77. T. kultiassovii M. Pop. ex S. Nikit. in Bot. Mat. Gerb. Bot. Inst. 

Akad. Nauk SSSR VII, 12 (1938) 265; Kovalevskaja in Fl. Uzb. VI, 431; 
Zakir. Fl. i Rast. Zeravsh. 2, 419. — T. ruber auct. fl. As. Med.: 
Kovalevskaja, op. cit. Plate XLIII, Fig. 3. 

Perennial. Root robust, vertical, cylindrical. Plants glaucous-green, 
10-30 cm high, usually branched from base. Stems thick, sulcate. Leaves 
broadly lanceolate, 10-15(20) mm wide in middle, usually recurved, 
undulate and scarious along margin, flocculose in leaf axils. Peduncles 
clavately thickened below capitula at fruiting, 4-6(8) mm thick. Capitula 
20-35 mm long at flowering. Involucral bracts eight, almost as long as 
florets; dry florets pinkish- violet. Achenes including pappus 3-6 cm and 
without pappus 1.8 cm long; body of achene about 1 cm long, finely 
sulcate and finely tuberculate, curved, gradually narrowed into about 0.8 
cm-long, thick beak; beak sulcate and finely tuberculate in lower half, 
not swollen at apex and without hairy annulus; pappus 1.8 cm long, as 
long as achene, light yellowish. Rowering April. 

Slopes of foothills. Soviet Central Asia: Syr-Darya, Pamiro-Alai 
(central Zeravshan, according to Zakirov). Endemic. Described from 
Fergana, vicinity of Santo. Type in Tashkent. 

Note. It differs from T. malicus S. Nikit. by having peduncles that 
are very strongly thickened at flowering, and also recurved and undulate 
leaves. 

78. T. kasahstanicus S. Nikit. in Bot. Mat. Gerb. Bot. Inst. Akad. 
Nauk SSSR VII, 12 (1937) 268.-7. rubrum auct. fl. turk. pro parte, non 
S.G. Gmel. 

Perennial. Plants grayish-green, up to 30 cm high. Stems branched 
below, with isolated flocculose flakes at leaf bases and below fertile 



196 

capitula. Middle leaves expanded at base up to 5-12(20) mm, 
semiamplexicaul, narrowed upward into narrowly linear or linear- 
lanceolate lamina, usually narrowly membranous along margin, sometimes 
sinuate and conduplicate. Peduncles more or less thickened below fertile 
capitula, (3)4-7 (sometimes up to 10) mm thick, fistular. Capitula about 
3 cm long at flowering, 4-5(6) cm long at fruiting. Involucral bracts 
eight, as long or almost as long as florets and achenes; florets pink-violet. 
Peripheral achenes straight or slightly curved, about 5 cm long, excluding 
pappus 2.5 cm long; body of achene 1.5 cm long, five-angled, with dense 
obtuse scales, sulcate above, gradually narrowed into somewhat thick, 
white, scabrous-sulcate and squamose, about 1 cm-long, basally smooth 
beak; beak almost one-third as long as achene, slightly swollen at apex; 
tomentose below pappus; pappus almost as long as achene, about 2.5 cm 
long, dirty white or yellowish; inner achenes not sculptured. Flowering 
May; fruiting June. (Plate VIII, Fig. 3.) 
196 Sandy deserts. — European Part: Lower Volga (Astrakhan, Lake 

Inder); Soviet Central Asia: Aralo-Caspian, Dzhungaria-Tarbagatai, 
Balkhash Region, Tien Shan, Syr-Darya. General distribution: Dzhungaria- 
Kashgaria. Described from sandy deserts in valley of Kurtsu River, a 
tributary of Hi River. Type in Leningrad. 

Economic Importance. A good food and fodder plant. 

Note. It differs from T. ruber S.G. Gmel. by having larger capitula, 
involucres almost equaling the florets and achenes, as well as peduncles 
that are thickened at fruiting. It differs from T. montanus S. Nikit. by 
having the middle cauline leaves expanded at the base and drawn out 
into a narrow lamina, a peduncle thickened below the capitula at fruiting, 
as well as by the pubescence and range. 

Section 17. Hebecarpus Boriss. in Addenda XXVIII, 725. — ^Peduncles 
thickened below capitula, fistular; involucral bracts eight, less often five, 
longer than or as long as florets; capitula large, 5-8 cm long; florets 
purple or violet; outer achenes straight, brownish, with straight, somewhat 
thick, smooth beak, gradually widening into squamose-tuberculate body 
of achene, almost as long beak; pappus more or less as long as achene. 
Large annuals or biennials. 

Type of section: T. porrifolius L. 

79. T. porrifolius L. Sp. pi. (1753) 789; DC. Prodr. VII, 112; Ldb. 
Fl. Ross. II, 2, 785; Boiss. Fl. or. Ill, 745; Schmalh. FI. II, 147 (T. 
parvifolium L. — sphalm.); Fedtsch. and Rer. Fl. Evrop. Ross. 1042. 

Annual or biennial. Root vertical, cylindrical, strong; root collar not 
covered with fibers. Stems simple or branched, glabrous or sparsely 



197 

pubescent, (30)60-125 cm high, thick. Leaves flat, linear-lanceolate, 
expanded at base, semiamplexicaul, straight, erect, acuminate. Peduncles 
thickened below capitula, tubular, fistular. Capitula 5-8 cm long, solitary, 
on long peduncles; involucral bracts eight, less often five, longer than or 
as long as florets, deflexed; florets purple or violet-red. Achenes straight, 
brownish; peripheral achenes squamose-tuberculate, gradually narrowed 
into straight, somewhat thick, smooth beak in broader body, apically 
extended into beak without thickening, almost as long as body of achene; 
pappus dirty white, more or less as long as achene with beak or slightly 
shorter, with hairy annulus at base. Flowering May to August. 

Cultivated for its edible roots, and sometimes growing wild. — 
European Part: Reported from Baltic Region (Lithuania, Latvia), Central 
Russian districts, Crimea and Lower Volga. General distribution: Central 
Europe, Atlantic Europe, Mediterranean Region, Balkans-Asia Minor. 

Note. The report of this species from the vicinity of Lake Inder 
probably relates to T. ruber s. 1. 



197 GENUS 1638. Geropogon L.'-2 

L. Sp. pi. ed. 2 (1763) 1109. 

Capitula less numerous, solitary, at apices of main stems, large, often 
numerous. Peduncle fistular, weakly thickened or not below capitula. 
Involucre of one row of similar or somewhat similar, usually remote bracts, 
3-4 cm long at fruiting, two times as long as florets; involucral bracts 
five to eight, straight, upward-spreading, linear or linear-lanceolate, 
herbaceous. Receptacle alveolate, ciliolate-ciliate from linear-subulate 
scales. Florets ligulate, pink or violet. Achenes straight, thin, roundish in 
cross section, scabrous, gradually narrowed into long straight beak as 
long as body of achene or two times as long, pappus half to two-thirds as 
long as achene; pappus different in different achenes: outer achenes with 
five erect, persistent, sharply scabrous, hard bristles; inner achenes with 
soft plumose deciduous bristles, without hairy annulus at base. Annuals 
or biennials with straight, usually unbranched stem, and narrowly linear, 
basally semiamplexicaul leaves. 

Type of genus: Geropogon hybridus (L.) Sch. Bip. 

A monotypic Mediterranean genus. 

1. G. hybridus (L.) Sch. Bip. in Webb, and Berth. Phyt. Canar. II 



' Treatment by A.G. Borissova. 

^ From the Greek words geron — old man, and pogon — ^beard; named on the basis of 
the white sericeous pubescence of the achene. 



198 

(1836) 411.— G. glabrum L. Sp. pi. ed. 2 (1763) 1109; Willd. Sp. pi. 3, 
1491, Pall. Ind. taur. 316; DC. Prodr. VIII, 111; Ldb. Fl. Ross. II 784; 
Boiss. Fl. or. Ill, 744; Grossh. Fl. Kavk. IV, 233; Kuthath. in Fl. Gruzii 
VIII, 636; Askerova in Fl. Azerb. VIII, 513. G. hirsutum L. Sp. pi. ed. 2 
(1763) 1109; Willd. 1. c.—G. austmlis Spreng. Syst. Veg. 3 (1826) 663.— 
Tragopogon hybridum L. Sp. pi. ed. I (1753) 789. — T. crocifolium auct. 
non L.: DC. Prodr. VII, 114, p. p.; Ldb. Fl. Ross. 3, 788, p. p.; Schmalh. 
Fl. II, 147.— r. hirsutus (L.) Tard. Ess. hist, natur. Bessar. (1841) 60.— 
Ic: Jacq. Hort. Vindob. t. 33. 

Annual. Root straight, virgate. Plants glabrous or weakly pubescent. 
Stems, 20-50(80) cm high, erect, usually simple, less often weakly 
branched, cylindrical. Leaves narrowly linear, long, erect, entire, 
acuminate, glabrous or pubescent at base, semiamplexicaul; lower leaves 
expanded at base. Peduncles hairy or glabrous below capitula, not or 
weakly thickened at fruiting, fistular. Capitula solitary, about 5 cm long, 

1 cm wide. Involucre almost two times as long as florets. Involucral bracts 
eight, less often five or six, narrow, linear or linear-lanceolate, acuminate, 

198 glabrous, 3-4 cm long at fruiting; florets pink or violet. Achenes including 
pappus about 5 cm long, without pappus 2.9-3.3 cm, almost roundish in 
cross section, linear-terete, straight, shallow sulcate, smooth below, sharply 
scabrous in middle and above, covered with rows of fine upward-spreading 
bristles; lower, broader body of achene gradually and imperceptibly 
narrowed into straight, stiff, rugose, about 1.5 cm-long beak; pappus of 
outer achenes of few subulate, sharp-toothed, scabrous, persistent, about 

2 cm long bristles; pappus of inner achenes comprising plumose, soft, 
deciduous hairs much shorter than achenes. Flowering April to June; 
fruiting May to June. (Plate VIII, Fig. 2.) 

Meadows of the lower mountain zone and plains, semideserts, among 
undergrowth in orchards, roadsides and fields. — European Part: Crimea, 
?Bessarabia; Caucasus: Eastern Transcaucasia, Talysh. General 
distribution: Mediterranean Region, Canary Islands, Madeira, Northern 
Africa. Described from Italy. Type in London. 

Subtribe 4. HYPOCHAERINAE Less. Synops. Compos. (1832) 130 
("Hypochaerideae"); DC. Prodr. VII, 1 (1838) 90; Benth. in Benth. and 
Hook. f. Gen. pi. II (1873) 168, 222, p. p.— Trib. Lactuceae sect. 1 
Archetypae Cass, in Diet. sc. nat. XXV (1822) 60, p. min. p.; sect. 2. 
Crepideae Cass. op. cit. 61, p. min. p.; sect. 4. Scorzonereae Cass. op. 
cit. 64, p. min. p. — Lactucees sect. 1 . Prototypes II. Urospermees Cass, 
in Diet. sc. nat. XL VIII (1827) 422; sect. 2. Crepidees Cass. Ibid., p. 
min. p.; Sect. 4. Scorzonerees Cass. Ibid., p. min. p. — Subtrib. 
Scorzonereae Dum. Fl. Belg. (1827) 63, p. p. — Trib. Cichorieae b) 
Scorzonereae Dum. Analyse fam. pi. (1829) 30, p. p. — Trib. Taraxaceae 



199 

D. Don in Edinb. New Philos. Journ. VI (1829) 307, p. p.— Trib. 
Hypochoerideae D. Don, Ibid, — Subtrib. Lampsaneae Less. op. cit. 126, 
p. min. p. — Subtrib. Hyoserideae Less. op. cit. 127, p. min. p. — Subtrib. 
Scorzonereae Less. op. cit. 131, p. p. — Sippen Hypochoerideae and 
Urospermeae Sch. Bip. in Flora XVII, 2 (1834) 475 and Picrideae and 
Leontodonteae Sch. Bip. op. cit. 476. — Subtrib. Rhagadioleae Benth. in 
Benth. and Hook. f. Gen. pi. II (1873) 168, 220, p. p. (solum gen. 
Rhagadiolus s. 1.). — Subtrib. Crepideae Benth. op. cit. p. min. p. — Subtrib. 
Scorzonereae Benth. Ibid. 169, 221, p. min. p. — Subtrib. Leontodontinae 
O. Hoffm. in Engl, and Prantl, Nat. Pflanzenfam. IV, 5 (1893) 36, p. p.; 
Stebbins in Madroiio XII, 3 (1953) 75.— Subtrib. Cichorinae O. Hoffm. 
op. cit. 353, p. min. p. — Pappus hairs of inner row plumose, outer setose, 
more or less toothed, or pappus entirely of plumose bristles; sometimes 
pappus of achenes in same capitulum different, coronate in some, of 
lanceolate-subulate scarious bristles in others; less often pappus entirely 
absent. Receptacle with membranous scales or smooth. Pollen grains 
spinose-cristate (echinolophate). Basic chromosome number x = 7, 6, 5, 
4, and 3. Plants usually with stiff scattered pubescence, hairs often forked. 
Type of subtribe: genus Hypochaeris L. 

199 GENUS 1639. Hypochaeris L.'-^ 

L. Sp. pi. (1753) 810. 

Capitula oblong-ovoid or almost campanulate, many-flowered, with 
involucre of imbricate bracts. Receptacle flat, with scarious, linear, long- 
acuminate scales. All florets ligulate, yellow. Achenes with or almost 
without beak, or only peripheral achenes without beak and inner ones 
beaked, smooth or scabrous; pappus two-rowed; bristles of outer row short, 
almost filiform, of inner row longer, plumose. Annual or biennial, less 
often perennial herbs with glabrous or hairy, simple or branched stem, or 
stemless. Leaves entire, toothed or pinnatilobate. 

Type of genus: Hypochaeris glabra L. 

About 15 species, predominantly Mediterranean. 

1. Involucral bracts glabrous, outer bracts oblong-oval, inner ones 
lanceolate-linear, thin, subobtuse; peripheral achenes without beak 
or with short beak 1. H. glabra L. 

+ Outer involucral bracts with narrow membranous border, ciliolate, 
lanceolate, obtuse; beak two to four times as long as achenes 
2. H. radicata. L. 



' Treatment by V.N. Vassilev. 

^ From the Greek words: ypo — for, and choiros — pigs; pigs relish the roots of this 



plant. 



200 

Section 1. Hypochaeris. — Sect. Euhypochaeris Duby, Bot. gall. I 
(1828) 306. — Outer achenes more or less without beak, inner ones with 
beak. 

Type of section: of genus species. 

1. H. glabra L. Sp. pi. (1753) 811; DC. Prodr. VII, 90; Ldb. Fl. 
Ross. II, 775; Boiss. Fl. or. III. 783; Sch. Bip. in Nova Acta Acad. Caes. 
Leopold. Carol. Nat. Cur. XXI, 1 (1845) 144. 

Annual. Root straight, digitate, simple or branched. Stems 15-30 cm 
high, 1-12 or more; outer ones ascending; middle ones erect, slender, 
cylindrical, sulcate, glabrous, branched, less often simple, with one 
capitulum, often in middle stems of rosula. Basal leaves 3-10 cm long, 
0.5-1.5 cm wide, often numerous, lanceolate, oblong, emarginate and 
200 runcinately pinnatipartite, acute, glabrous, or pubescent above and along 
margin. Capitula always erect, small, 1.5-2 cm long, 1.5-2 cm wide. 
Involucre almost cylindrical, imbricate, glabrous, slightly shorter than 
achenes with pappus. Outer involucral bracts oblong-ovate; inner ones 
lanceolate-linear, thin, subobtuse, somewhat dark above. Receptacle 
squamose; scales slightly shorter than pappus, long-acuminate, one-veined; 
inner florets golden-or lemon-yellow, with filiform tube longer than five- 
toothed ligule; peripheral achenes without beak or with a short beak, inner 
achenes long-beaked; all achenes with 15 well-developed, dark brown 
furrows, as also beak, scabrous; pappus dirty white, of two rows of bristles; 
outer bristles short, scabrous, scarcely plumose; inner bristles ten, longer, 
plumose. Flowering June to July; fruiting July to August. 

Sands, as a weed in fields and pastures. — European Part: Baltic 
Region, Upper Dnieper, Middle Dnieper, Upper Dniester. General 
distribution: Scandinavia, central Europe, Atlantic Europe, Mediterranean, 
Balkans-Asia Minor, North America (western states, introduced), Australia 
(introduced). Described from western Europe. Type in London. 

Section 2. Porcellites (Cass.) DC. Prodr. VII (1838) 91.— Porcellites 
Cass, in Diet. sc. nat. XXV (1822) 64 and XLIII (1826) 42, p. p. pro 
gen. Achenes with beaks of equal length. 

Type of section: Hypochaeris radicata L. 

2. H. radicata L. Sp. pi. (1753) 811; DC. Prodr. VII, 91; Ldb. Fl. 
Ross. II, 776; Boiss. Fl. or. Ill, 783; Sch. Bip. in Nova Acta Acad. 
Caes, Leopold. -Carol. Nat. Cur. I (1845) 155. — Achyrophorus radicatus 
(L.) Scop. Fl. carniol. ed. 2, II (1772) 117. — Porcellites radicata 
(L.) Cass, in Diet. sc. nat. XLIII (1826) 43. — Hypochaeris runcinata 
Gilib. Fl. Lithuan. Ill (1785) 226.— Ic: Gaertn. De fruct. II (1791) t. 
159. 



201 

Perennial. Root virgate, fusiform. Stem erect, 25-60 cm high, 
glabrous, usually branched, with very small sessile leaves, usually with 
few capitula at apices of branches. Leaves of basal rosette oblong- 
lanceolate, toothed, lyrate or runcinately pinnatipartite, glabrous or hispid 
on both sides. Capitula large, 2.5-3.0 cm long, up to 2 cm wide at 
flowering and fruiting. Involucre imbricate, almost half as long as florets; 
outer involucral bracts lanceolate, obtuse, with narrow membranous border 
and ciliolate; inner bracts narrowly lanceolate, with long cusp; all 
involucral bracts absolutely glabrous or with long stiff cilia-like hairs 
along margin, or also with long ferruginous hairs on dorsal side. Corolla 
yellow with ligule almost two times as long as tube, with five linear 
oblong small teeth at apex. Achenes with beak, beck two to four times as 
long as achene, sulcate, transversely striate, prickly above and along beak; 
bristles of outer row of pappus short, simple, those of inner row two 
201 times as long as outer, plumose. Flowering, second half of June to July; 
fruiting July to August (Plate XII, Fig. 6). 

Mountain and plains meadows, forest glades, pine forests. — European 
Part: Ladoga-Ilmen, Upper Volga, Baltic Region, Upper Dnieper, Middle 
Dnieper, Upper Dniester, Bessarabia, Black Sea Region, Lower Don; 
Caucasus: Ciscaucasia, Dagestan, Eastern and Western Transcaucasia. 
General distribution: Scandinavia, central Europe, Atlantic Europe, 
Mediterranean, Balkans-Asia Minor, North America, Australia 
(introduced). Described from western Europe. Type in London. 



GENUS 1640. Achyrophorus Scop. 



1.2 



Scop. Fl. carinol. ed. 2, II (1772) 116. 

Capitula one to three, large, ovoid, often hemispherical or 
campanulate, with involucre of imbricate bracts. Receptacle flat, with long, 
membranous, conduplicate, linear, acuminate scales enclosing florets at 
base. All florets ligulate, yellow. Achenes with or without beak, terete- 
sulcate, oblong; pappus one-rowed, of plumose bristles. Perennial herbs 
with simple or weakly branched, more or less leafy stem, and rosette of 
large basal leaves. 

Type of genus: Achyrophorus maculatus (L.) Scop. 

About 20 species distributed in Europe, Asia, and mainly in South 
America. 

1. Outer involucral bracts linear-lanceolate, dorsally hispid; stem 
branched, with few capitula 1. A. maculatus (L.) Scop. 



' Treatment by V.N. Vassilev. 

- From the Greek words: achyron — straw, and pharien — to bear. 



202 

+ Outer involucral bracts ovate or oblong-ovate, ciliate along margin, 
dorsally with crisped hairs; stem simple, one-headed 2. 

2. Cauline leaves amplexicaul, capitula up to 2.5 cm long and wide; 

outer involucral bracts ciliate along margin 

2. A. ciliatus (Thunb.) Sep. Bip. 

+ Cauline leaves narrowed toward base; capitula up to 6 cm wide 
and 1 cm long, tomentose with upper part of stem; outer involucral 

bracts ciliate along margin, dorsally with long crisped hairs 

3. A. uniflorus (Vill.) Bluff, and Fingerh. 

Section 1. Achyrophorus. — Sect. Phanoderis DC. Prodr. VII (1838) 
92. — All achenes distinctly narrowed into long beak; beak of outer achenes 
shorter than that of inner achenes. 

Type of section: Achyrophorus maculatus (L.) Scop. 

202 1. A. maculatus (L.) Scop. Fl. Carinol. II (1772) 116; DC. Prodr. 

VII. 92.— Ldb. Fl. Ross. II, 776; Kryl. Fl. Zap. Sib. XI, 2993.— 
Hypochaeris maculata L. Sp. pi. (1753) 810; Pall. Reise III, 687; Ldb. 
Fl. Alt. IV, 164; Kryl. Fl. Alt. Ill, 737.: Ic: Garcke, 111. Fl. Deutschl. 
(1895) 354. 

Perennial. Root vertical, up to 1.5 cm thick near root collar. Stem 
erect, 30-120 cm high, densely (especially below) covered with stiff 
spreading hairs, less often almost glabrous, with one, less often two or 
three small, narrow leaves, simple or branched, covered with dark brown 
scales at base. Basal leaves oblong-elliptical, usually obtuse, narrowed 
toward base, sinuate-toothed, less often almost entire, often with reddish- 
brown spots, with scattered white, stiff, appressed hairs on both sides, 
less often one or other side glabrous, 5-25 cm long, 1.5-7 cm wide; 
cauline leaves sessile, oblong or lanceolate, acuminate. Capitula solitary 
at apices of main stem or lateral branches with oblong involucre, 15-30 
mm wide, 15-20 mm long; outer bracts linear-lanceolate, dorsally hispid, 
inner; ones linear-oblong, glabrous or dorsally floccose, more or less 
membranous; all bracts yellowish-tomentose along margin. Ligules yellow, 
outer ones one and one-half times or more as long as involucre; scale 
acuminate, pubescent on outer side and along margin, slightly shorter 
than corolla or achene with pappus. Achenes yellowish-brown, with five 
longitudinal ribs, transversely rugose, with long beak, almost as long as 
achene; pappus one-rowed, of plumose hairs. Flowering June to July; 
fruiting June to August. (Plate XII, Fig. 4.) 

Forest and steppe, plains and mountain meadows, forest edges, open 
birch and pine forests. — European Part: Dvina-Pechora, Ladoga-Ilmen, 
Upper Volga, Baltic Region, Volga-Kama, Upper Dnieper, Upper Dniester, 
Middle Dnieper, Volga-Don, Trans- Volga Region, Bessarabia, Black Sea 



203 

Region, Lower Don, Lower Volga Region; Caucasus: Ciscaucasia, Western 
Transcaucasia; Western Siberia: Ob', Upper Tobol, Irtysh, Altai; Eastern 
Siberia: Angara-Sayans, Dauria (west). General distribution: Scandinavia 
(southern part), central and Atlantic Europe, Mediterranean Region 
(northern Italy), Balkans-Asia Minor (Balkans). Described from western 
Europe. Type in London. 

Section 2. OreophUa (D. Don) DC. Prodr. VII (1838) 93.— Oreophila 
D. Don. in Trans. Linn. Soc. XVI (1830) 178, pro gen. — All achenes 
with short beak, without beak, or with long beak; outer involucral bracts 
ovate or oblong-ovate, ciliate along margin. 

Type of section: Achyrophorus sessiliflorus (Kunth) V. Vassil. comb, n 

2. A. ciliatus (Thunb.) Sch. Bip. in Nova Acta Acad. Caes. Leopold. 
203 Carol. Nat. Cur. XXI, I (1845) 128; Kryl. Fl. Zap. Sib. XI, 299A.— Arnica 
ciliata Thunb. Fl. Jap. (1784) 318. — Hypochaeris grandiflora Ldb. Fl. 
Alt. IV (1833) 164; Kryl. Fl. Alt. Ill, 738; Kom. Fl. Manchzh. Ill, 764.— 
Achyrophorus aurantiacus DC. Prodr. VII (1838) 93.— A. grandiflorus 
(Ldb.) Ldb. Fl. Ross. II (1844-1846) 777; Maxim. Prim. Fl. amur. 176.— 
Hypochaeris ciliata (Thunb.) Makino in Bot. Mag. Tokyo XXII (1908) 
37.—//. aurantiaca Turcz. ex DC. Prodr. VII (1838) 94 in syn.— 
Amblachaenium aurantiacum Turcz. ex DC. op. cit. 94 in syn. — 
Oreophila sibirica C.A. Mey. ex Turcz. in Bull. Soc. Nat. Mosc. XI (1838) 
95.— Ic: Ldb. Ic. pi. fl. ross., t. 440. 

Perennial. Root straight, stiff. Stem straight, 30-65 cm high, simple, 
with dry brown scales at base, finely sulcate, rather densely hispid 
throughout or only below, less often almost entire stem glabrous. Basal 
leaves in rosettes, petiolate, oblong, sUghtly expanded in upper part, acute, 
sharp-toothed, setose-ciliate; lower cauline leaves like basal; middle and 
upper leaves sessile, amplexicaul, reduced toward apex, oblong or 
elliptical, finely toothed, hispid beneath. Capitula large, with almost 
hemispherical involucre, about 2,5 cm long and wide. Outer bracts ovate 
or oblong-ovate, ciliate along margin; inner bracts lanceolate, acute. 
Ligules bright orange, with narrow tube, slightly longer than limb; limb 
margin with five cristate denticles. Achenes terete, slightly narrowed 
upward; pappus one-rowed, of plumose hairs. Flowering June to July; 
fruiting end of July to beginning of August. 

Bottomland and mountain meadows, less often in thickets of shrubs, 
in drier habitats, especially sandy soils. — Western Siberia: Altai; Eastern 
Siberia: Lena-Kolyma, Dauria; Far East: Zeya-Bureya, Ussuri. General 
distribution: Mongolia, northeastern China, northern part of Korean 
Peninsula; in Japan, only under cultivation as an ornamental plant. 
Described from Japan. Type in Uppsala. 



204 

Economic Importance. Can be recommended as an ornamental plant. 

3. A. uniflorus (Vill.) Bluff, and Fingerh. Compend. Fl. Germ. II 
(1825) 301.— Hypochaeris uniflora Vill. Prosp. pi. Dauph. (1779) 37.— 
H. foliosa Vill. Fl. Dauph. Ill (1785) 25.— H. helvtica Wulf. in Jacq. 
Miscell. Austr. Ill (1785) 25. — Achyrophorus helveticus (Wulf.) Homem. 
Hort. hafn. II (1815) 772; Less. Synops. compos. (1832) 130.— Porcellites 
helvetica (Wulf.) Cass, in Diet. sc. nat. XLVIII (1826) 43.— Ic: Jacq'. 
Ic. pi. rar. (1781) t. 165; Vill. op. cit. (1789) 61, t. 23. 

Perennial. Root cylindrical, fusiform. Stem solitary (very rarely two), 
erect, 10-40 cm high, strong, ribbed-sulcate, thickened above, covered 
with simple short fibers. Basal leaves oblong-lanceolate, 8-20 cm long, 
up to 2 cm wide, sinuate-toothed, acute, hispid, on short, less often on 
204 long petioles; cauline leaves two or three, sessile, smaller than basal, often 
very small, narrow leaves present in upper part of stem, resembling outer 
involucral bracts. Capitulum one, at stem apex, up to 6 cm wide and 4 
cm long at flowering. Involucre imbricate, tomentose below, like upper 
part of stem; outer and middle involucral bracts ovate or oblong-ovate, 
ciliate along margin, with simple crisped, somewhat long, light gray hairs 
on dorsal side; inner bracts narrowly lanceolate. Receptacles squamose; 
scales linear, one-veined, long-acuminate, as long as achene with pappus. 
Ligules golden, peripheral twice as long as involucre and twice as long 
as tube. Achenes yellowish; peripheral ones harder and shorter, inner ones 
thinner and longer; usually 3 cm long including pappus, 2 cm without 
pappus, with longitudinal furrows; pappus persistent, of 22-25 plumose 
whitish bristles, one-rowed. Flowering June to July; fruiting July to August. 

Alpine zone. — European Part: Upper Dniester (Carpathians). General 
distribution: central and western Europe. Described from western Europe. 
Type in Paris. 

Economic Importance. Suitable for cultivation as an ornamental 
plant. 

GENUS 1641. Leontodon' 2 

L. Sp. pi. (1753) 798. 

Capitula many-flowered; involucre more or less imbricate, one- to 
three-rowed, outer bracts shorter. Receptacle without scales, alveolate, 
fibrous or finely punctate, with small scaly hairs along edge of pits. All 



' Treatment by V.N. Vassilev. 

^Name derived by combining two latin words: leo — lion, and dens — tooth; named 
on the basis of the leaf shape resembling lion teeth. 



205 




205 Plate XH 

1 — Leontodon croceus Haenke.; 2 — L gutzulorum V. Vassil.; 3 — L. rilaensis Hayek.; 

4 — Achyrophorus maculatus (L.) Scop.; 5 — Urospermum picroides (L.) Desf.; 

6 — Hypochaeris radicata L.; 7 — Helminthia echioides (L.) Gaertn. 



206 

florets ligulate, yellow or orange. Achenes terete, more or less fusiform, 
without or with short beak, longitudinally sulcate, transversely rugose; 
pappus persistent, one- or two-rowed, its bristles not connate at base, 
bristles of inner row plumose, of outer row always shorter, often with 
simple teeth. Stem simple, one-headed, or branched and with few capitula. 
Leaves in basal rosette, toothed or pinnatifid, less often subentire. Annual 
or perennial herbs. 

Type of genus: Leontodon hispidus L. 

206 As many as 45 species have been recognised in the genus, many of 

which are distributed in the Mediterranean Region. To the east, the genus 

reaches up to the western part of western Siberia; in the north it reaches 

the tundra zone. A few species are present in North and South America. 

1. Plant covered with simple hairs or absolutely glabrous 2. 

+ Plant covered with forked or stiff stellate hairs or simple hairs 

mixed with forked hairs 10. 

2. Plants absolutely glabrous 6. L. danubialis Jacq. 

+ Plants never absolutely glabrous 3. 

3. Rhizome creeping, horizontal; leaves and stem covered with sparse 
and scattered long silky hairs 7. L. repens Schur. 

+ Rhizomes short, as if truncate 4. 

4. Scape with one capitulum, less often with two 5. 

+ Scape always with two or more capitula 8. 

5. Leaves entire, glabrous, smooth; scape up to 10 cm high, as also 
involucral bracts, sparsely pubescent 3. L. rilaensis Hayek. 

+ Leaves toothed or more or less divided 6. 

6. Plant short, 5-15(18) cm high, scape with one, less often with 
two capitula 2. L. pseudotaraxaci Schur. 

+ Plant taller, 15-35 cm high, always one-headed 7. 

7. Florets orange; achenes without beak; pappus one-rowed 

9. L. croceus Haenke^ 

+ Florets yellow; achenes with short beak; pappus two-rowed 

8. L. schischkinii V. Vassil. 

8. Capitula densely covered with long stiff rusty-greenish hairs; 
pappus two-rowed 4. L. keretinus Nyl. 

+ Pubescence of capitula different; pappus one-rowed 9. 

9. Involucre with white arachnoid pubescence (f. autumnalis) or, in 
addition, with long, setose, rusty, less often almost white or black 
hairs (f. pratense); achenes fusiform 1. L. autumnalis L. 

+ Involucre densely covered with long, dark, soft hairs; achenes 

terete, beaked 5. L. gutzulorum V. Vassil. 

10. Root long, radish-shaped, merging with rhizome above; scape 
simple or with one or two branches; outer involucral bracts ciliate; 



207 

bristles white, short, forked; achenes with long beak 11. 

207 + Rhizome short, as if truncate; scape always one capitulum; outer 
involucral bracts without forked bristles; achenes with short beak 
13. 

11. Pubescence on leaves and stem of gray, very short, forked and 
stellate hairs, hence whole plant grayish, as if velutinous 

12 L. kotshyi Boiss. 

+ Pubescence on leaves and stem of long gray forked hairs, hence 
plant setose-hairy 12. 

12. Capitula 2-2.5(3) cm long, narrowly obconical before flowering, 
almost cylindrical; achenes 15-20 mm long 

11. L. asperrimus (Willd.) Boiss. 

+ Capitula shorter, up to 1.5 cm long, obconical before flowering; 

achenes 8-12 nmi long 

10. L. asperus (Waldst. and Kit.) Boiss. 

13. Stem glabrous; leaves and involucral bracts glabrous or sparsely 
hairy; hairs on leaves simple and forked 

14. L. caucasicus (M.B.) Fisch. 

+ Stem scabrous from short forked hairs; leaves rather densely 
covered with short or long, forked hairs; involucre densely covered 
with simple setose hairs 13. L. hispidus L. 

Subgenus 1. Scorzoneroides (Moench) V. Vassil. comb. nova. — 
Scorzoneroides Moench, Meth. pi. (1794) 549, pro gen. — Apargia Scop. 
Fl. Camiol. ed. 2, II (1772) 113, p. p. — Subgen. Euleontodon Widder in 
Oesterr. Bot. Zeitschr. (1931) 146. — Plants with simple hairs or glabrous; 
rhizome reduced, very rarely (in L. repens) creeping; scapes simple or 
more or less branched, thickened below capitula. 

Type of subgenus: Leontodon autumnalis L. 

Series 1. Autumnales V. Vassil. — Leaves and stem glabrous or weakly 
pubescent; stem ascending; achenes not beaked; pappus one-rowed. 

Leontodon montanus Lam. and L. pyrenaicus. Gouan, from outside 
of the limits of our flora, also belong to this series. 

1. L. autumnalis L. Sp. pi. (1753) 798; Ldb. Fl. Ross. II, 778; 
Boiss. Fl. or. Ill, 729. — Scorzoneroides, Autumnalis (L.) Moench, Meth. 
pi. (1794) 549. — Hedypnois autumnalis (L.) Huds. Fl. Angl. ed. 2 (1762) 
341. — Picris autumnalis (L.) All. Fl. Pedem. 1 (1785) 210. — Apargia 
autumnalis (L.) Hoffm., Deutschl. Fl. ed. II (1804) 113. — Ic: Fedtsch. 
and Fler. Fl. Evrop. Rossii, Fig. 1062. 

Perennial. Rhizomes reduced, as if truncate, with numerous stiff hairs. 
Stems usually branched, less often simple. Plants glabrous or more or 



208 

less densely covered with simple hairs (f. pratense Koch). Basal leaves 

208 prostrate, elongated, rather wide, with long-winged petioles, toothed, 
pinnatifid or pinnatipartite with linear lobes, less often entire or almost 
entire (f. subintegrifolius m.). Peduncles long, fistular, thickened at apex, 
with linear or subulate, appressed leaflets. Capitula erect before flowering, 
up to 2 cm long at flowering, 2.5 cm wide at fruiting. Involucre of 
lanceolate bracts with white arachnoid pubescence (f. autumnalis m.) or, 
also with long setose ferruginous, rarely almost white or black, hairs (f. 
pratensis Koch). Corolla yellow, very rarely pale violet. Achenes fusiform, 
5 mm long, slightly narrowed at both ends, sulcate, transversely rugose; 
pappus up to 6-7 mm long, dirty white, one-rowed, of plumose, barbed 
bristles, weakly thickened toward base; less often simple barbed bristles 
found among plumose ones. Flowering July to October; fruiting August 
to October. 

Fields, pastures, meadows, shrubby thickets, open forests. — Arctic 
Region: Arctic Europe (Kola Peninsula); European Part: Karelia-Lapland, 
Dvina-Pechora, Ladoga-Ilmen, Upper Volga, Volga-Kama, Upper Dnieper, 
Middle Dnieper. Volga-Don, Trans- Volga Region, Bessarabia, Black Sea 
Region, Crimea, Lower Don, Lower Volga; Caucasus: Ciscaucasia, 
Western and Southern Transcaucasia; Western Siberia: Ob', Upper Tobol, 
Irtysh, Altai; Eastern Siberia: Angara-Sayans (village of Mysovka, 
introduced); Far East: Kamchatka (introduced), Ussuri (introduced), 
Sakhalin (introduced). General distribution: Scandinavia, central Europe, 
Atlantic Europe. Described from northern Europe. Type in London. 

Note. This species is extremely polymorphic, but there is little 
justification for elevating its forms to the .rank of species. 

2. L. pseudotaraxaci Schur. Enum. pi. Transsilv. (1866) 357. — L. 
taraxaci var. tatricus Kot. Distr. pi. in mont. Tatr. (1890) 356. — L. clavatus 
Sag. and G. Schm. Fl. Carp, centr. II (1891) 254.— L. tatricus (Kot.) 
Wolocz. Fl. Polon. exs. (1897) 545. — Ic: Javorka and Csapody, Icon. 
Fl. Hung. (1933) 554, No. 3984. 

Perennial. Rhizomes decurrent, many-headed; scape ascending, with 
one, less often two capitula, 8-15 cm long, distinctly thickened in upper 
part, sulcate, densely covered with simple, thick, soft hairs and scaly leaves 
above merging with involucral bracts, glabrous or almost glabrous below. 
Basal leaves oblong-spatulate, obtuse, narrowed into long petiole, 
runcinately pinnatifid, with 3-15 cm long petiole, sometimes with three 
unequal, almost triangular, sharp teeth in middle on both sides, glabrous 
or with less numerous, long, simple hairs beneath. Capitula turbinate, 
drooping before flowering; involucral bracts linear, dorsally covered with 
blackish hairs, their tips and margin white-sericeous. Corolla orange, tube 

209 scatteredly white-sericeous; ligules truncate above, five-toothed. Pappus 



209 

dirty white, weakly ferruginous on drying; pappus bristles plumose, one- 
rowed, strongly thickened at base; achenes flat, somewhat curved, sulcate, 
smooth, matty. Flowering August; fruiting August to September. (Plate 
XIII, Fig. 1.) 

Limestone rocks of alpine zone, swales. — European Part: Upper 
Dniester. General distribution: Central Europe (Transylvanian Alps). 
Described from Kronstadt-Alps. Type in Lvov. 

3. L. rilaensis Hayek in Denkschr. Akad. Wissensch. 94 (1918) 198. 
Widder in Oesterr. Bot. Zeitschr. LXXVI (1927) 293.— L. pyrenaicus, d. 
integerrimus, Schur. Enum. PI. Transsilv. (1866) 358. — Ic: Javorka and 
Csapody, Icon Fl. Hung. (1933) 554, No. 3983a. 

Perennial. Rhizome more or less obliquely truncate; scape usually 
solitary, erect, 10-15 cm high slender, two times as long as rosulate leaves, 
leafless below, glabrous, thickened above, leafless or with linear-lanceolate 
scaly leaves, especially below capitula in thickened part, and covered 
with white-arachnoid pubescence. Leaves 3-6 cm long, spatulate-linear 
or lanceolate, narrowed toward base, with winged petioles, entire or, very 
rarely, with one to three indistinct teeth, glabrous. Capitula always erect. 
Involucral bracts linear-lanceolate, loosely imbricate; outer bracts with 
blackish or light green hairs on dorsal surface (f. viridulus m.), white- 
arachnoid along margin, inner bracts glabrous, translucent; receptacle 
glabrous; ligules yellow. Achenes terete-fusiform, sulcate, sometimes also 
weakly transversely rugose; pappus plumose, very rarely at apex with small 
simple bristles, lustrous, dirty white. Flowering July. (Plate XII, Fig. 3). 

Pastures in subalpine zone. — European Parti Upper Dniester. General 
distribution: central Europe (Carpathians), Balkans-Asia Minor (Balkan 
Peninsula). Described from the Balkan Mountains. Type in Vienna. 

Note. Our material differs from the type specimen by the absence of 
blackish hairs on the upper, expanded part of the scape and by having 
light green hairs on the dorsal surface of the outer involucral bracts. 
Apparently, L. pyrenaicus Gouan is the closest species to L. rilaensis. 

Series 2. Keretini V. Vassil. — Stem branched, ascending; capitula 
rather densely covered with long, relatively soft hairs; achenes short- 
beaked; pappus one-rowed or two-rowed. 

4. L. keretinus Nyl. Spicil. pi. fenn. I (1843) 24; Ldb Fl. Ross. II, 
779. 

Perennial. Rhizome as if truncate, with numerous fibrous roots, many- 
headed, producing few slightly arcuate stems. Stems sulcate, glabrous up 
210 to branching, weakly pubescent and branched above, 15-45 cm high, 
with several capitula. Basal leaves glabrous or with sparse simple hairs, 
smooth, oblong-lanceolate, lamina decurrent on petiole, 3-18 cm long. 



210 

deeply pinnatifid, with narrowly lanceolate or linear lobes, usually antrorse; 
usually leaves simple, linear, sometimes few large pinnatifid leaves near 
fork. Peduncles distinctly thickened, with small linear leaves, gradually 
merging with involucral bracts. Capitula turbinate; involucral bracts linear, 
acute, densely covered with simple, long, somewhat stiff, ferruginous hairs, 
with soft, short, silky, dirty white hairs above; Ugules orange. Achenes 
5-7 mm long, brown, sulcate, transversely rugose, with short beak; pappus 
5-7 mm long, dirty white, plumose, two-rowed; pappus bristles almost 
not thickened at base, outer bristles short. Flowering June to July; fruiting 
July to August. 

River valleys, crop fields, pastures. — European Part: Karelia-Lapland, 
Dvina-Pechora, Ladoga-Ilmen, Upper Volga, Volga-Kama. General 
distribution: Scandinavia (Finland). Described from Keret River. Type in 
Leningrad. 

Note. The small number of specimens was collected from the northern 
and northwestern part of Eastern Europe. In general appearance, it is 
sharply distinguished not only from typical, L. autumnalis L., but also 
from its var. pratensis Koch, from which it differs by the brighter color 
of its corolla and the nature of the pubescence on the capitula and 
receptacle. 

5. L. gutzulorum V. Vassil. in Bot. Mat. Gerb, Bot. Inst. Akad. 
Nauk SSSR, XXI (1961) 397. 

Perennial. Rhizome short, as if truncate, with numerous fibrous roots, 
many-headed. Stems 18-25 cm high, arcuately ascending, branched, with 
two to four capitula, glabrous, smooth, distinctly thickened above and 
covered with dark soft hairs, with small subulate acuminate leaves near 
fork and above. Basal leaves glabrous, smooth, sinuate-long-toothed or 
pinnatifid, with antrorse lobes, somewhat acuminate, less often leaves 
almost entire, 7-10 cm long, 1-1.5(2) cm wide. Capitula drooping before 
flowering, erect during and after flowering, turbinate, 2 cm long at 
flowering, 2 cm wide. Involucral bracts dark green, densely covered with 
dark, soft, long hairs, with dirty-white short silky hairs at apex and along 
margin; corolla yellow, much longer than involucre. Achenes yellowish, 
terete, sulcate, narrowed into beak, transversely rugose, 7 mm long, 0.5 
mm wide; pappus one-rowed, up to 7 mm long, somewhat sordid; its 
bristles plumose, somewhat thickened at base. Flowering July; fruiting 
August. (Plate XII, Fig. 2.) 

Alpine patches of meadow. — European Part: Upper Dniester (Goverla 

Mountain). General distribution: central Europe. Described from Goverla 

Mountain. Type in Leningrad. 

211 Note. Extremely close to L pseudotaraxaci Schur, on the basis of 

the pubescence on the peduncles and involucral bracts, but it is well 



211 

differentiated by the branched stem, glabrous leaves and large size of the 
entire plant. 

Series 3. Danubiales V. Vassil. — Scape always one-headed, glabrous 
or weakly pubescent; achenes without beak; pappus one-headed, except 
in L. croceus Haenke. 

6. L. danubialis Jacq. Enum. Stirp. vindobon. (1762) 139, 270. — L. 
hastilis L. Sp. pi. ed. 2 (1763) 1123; Ldb. Fl. Ross. II, ISO.— Apargia 
dubia Willd. Sp. pi. ed. IV, III, 3 (1803) 1549.— Leontodon hyposeroides 
var. glaberrimmus Schur, Enum. pi. Transsilv. (1866) 359. — L. clavatus 
Schur, Phyt. Mitt. (1876) 193.— P/cm danubialis (Jacq.) All. Fl. Pedem. 
I (1785) 211.— Ic: All. Ibid. t. 70, fig. 3; Jacq. Fl. Austr. II (1774) t. 
164; Javorka and Csapody, Ic. Fl. Hung. (1929) 555, No. 3981. 

Perennial. Rhizome truncate, short, with numerous fibrous roots; 
scapes 15-^0 cm high, glabrous, smooth, one or two from single rhizome, 
each one-headed, sometimes reddish at base, entirely leafless or with one 
or two linear scaly leaves above, very rarely with one or two scaly leaves 
in lower part of scape, usually not thickened below capitulum. Leaves of 
rosette spamlate, toothed, sinuate-toothed or emarginate-toothed, less often 
almost entire, thin, glabrous, on rather long winged petioles, 3-22 cm 
long (including petioles), 0.9-3.0 cm wide. Capitulum erect; involucral 
bracts dark green, imbricate, linear, subulate, glabrous, ciliate only along 
margin, cilia white, short; outer bracts deflexed; ligules yellow, with long 
white hairs at base. Achenes 6 mm long, brownish, slightly curved, 
longitudinally sulcate, transversely rugose, without beak; pappus 8 mm 
long, two-rowed, setose, bristles of outer row thin, short, simple, barbed, 
inner bristles plumose, strongly thickened toward base. Flowering July. 

Meadows, borders of fields, roadsides, glades in beech forest zone. — 
European Part: Ladoga-Ilmen, Upper Dnieper, Middle Dnieper, Upper 
Dniester, Black Sea Region, Crimea; Caucasus: Ciscaucasia, Dagestan, 
Eastern, Western, and Southern Transcaucasia, Talysh. General 
distribution., central Europe, Atlantic Europe, Balkans-Asia minor (north 
of Balkan Peninsula). Described from vicinity of Vienna. Type in Vienna. 

Note. In the almost complete absence of pubescence and shape of 
the leaves, this species is closest to L. montaniformis Widd. and L. 
pseudotaraxaci Schur, 

7. L. repens Schur in Verb. Siebenb. Ver. X (1859) 138, 148, 173; 
Degen in Mag. Bot. Lapok 23 (1924) 91-94. — L. hispidus b. caucasicus 
GrSesku, Consp. Fl. Roman. (1898) 353;- Unger. Fl. Siebenburg. (1925) 
498.— L. kulczinskii M. Pop. Ocherk Rast. i Fl. Karpat (1949) 299.— 

212 Ic: M. Pop. op. cit. 255; Javorka and Csapody, Ic: Fl. Hung. I (1929) 
554, No. 3980a. 



212 

Perennial. Rhizome horizontal, rather long, creeping or obliquely 
ascending, branched, developing rosette; scapes (stem) one-headed, 25- 
45 cm, high, less often shorter, with rosette of basal leaves, leafless above 
or with a small leaf below capitulum, sulcate, covered with simple scattered 
hairs, thickened above below capitulum, densely hairy; leaves narrow, 
oblong-lanceolate, narrowed toward base, 9-23 cm long, 1-3 cm wide, 
finely toothed or almost entire, dark green, with dense simple hairs on 
both sides along margin and on lower side along midrib. Capitula blackish, 
with involucre of appressed bracts, outermost bracts loose, slightly 
deflexed, narrowly lanceolate, distinctly narrowed above, weakly 
translucent along margin, smooth, with long whitish hairs along dorsal 
surface, obtuse-ciliate above; ligulate florets yellow, often brownish on 
outer side, two times as long as involucral bracts, toothed above, with 
blackish teeth. Pappus 9 mm long, dirty white, of unequal bristles, inner 
bristles plumose, outer ones short, scabrous; achenes chestnut-colored, 5 
nrni long, without beak. Flowering July, August; fruiting August to 
September. (Plate XIII, Fig. 5.) 

Meadows of subalpine and alpine zones. European Part: Upper 
Dniester. General distribution: Central Europe (Transylvanian Alps). 
Described from Transylvania. Type in Vienna. 

8. L. schischkinii V. Vassil. in Bot. Mat. Gerb. Bot. Inst. Akad. 
Nauk SSSR XXI (1961) 398. 

Perennial. Rhizome truncate, with numerous stiff hairs, one-, two- or 
three-headed; stem scape, 15-35 cm high, one-headed, weakly thickened 
above, smooth, only near apex and at base with very short, appressed, 
white, silky hairs and one or two linear-lanceolate or linear-subulate small 
leaves (or without them). Basal leaves 5-10 cm long, 1-2 cm wide, 
spatulate, glabrous or weakly pubescent with simple hairs, remotely 
sinuate-toothed or sinuate-incised, gradually narrowed into winged petiole. 
Capitula drooping before flowering, erect at flowering, 2.5 cm long, 3.5 
cm wide (at flowering). Outer involucral bracts short, linear, deflexed; 
inner ones narrowly lanceolate, acuminate, white-sericeous along margin, 
214 with similar fine hairs on dorsal side; outer most involucral bracts glabrous, 
with scarious margin; receptacle glabrous. Corolla yellow; ligules 
with five very short teeth, darker than remaining part of ligule, with 
long, crisped, white, silky hairs at base. Achenes brown, fusiform, 
sulcate, transversely rugose, somewhat curved, up to 0.5 cm long; pappus 
up to 0.8 cm long, dirty white, two-rowed, of plumose bristles; bristles of 
outer row much shorter and thinner than inner, inner bristles strongly 
thickened toward base. Flowering July; fruiting August. (Plate XIII, 
Fig. 3.) 

Glades, forest edges, mountain slopes. — European Part: Upper 



213 




213 Plate Xffl 

1 — Leotodon pseudotaraxaci Schur; 2 — L hispidus L.; 3 — L schischkinii V. Vassil.; 

4 — L. asperrimus (Willd.) Boiss.; 5 — L. repens Schur; 6 — L asperus (Waldst. and Kit.) 

Boiss.; 7 — L. Kotschyi Boiss. 



214 

Dniester. General distribution: central Europe. Described from eastern 
Carpathians. Type in Leningrad. 

Note. It differs from L pyrenaicus Gouan, with which it was confused, 
by having larger and more deeply divided or boldly dentate leaves and 
involucral bracts with fine white arachnoid pubescence, whereas the 
pubescence of the involucral bracts in L. pyrenaicus consists of dark, 
sometimes almost black, long hairs. 

9. L. croceus Haenke in Jacq. Collectanea II (1788) 16; Rchb. Fl. 
Germ. exc. (1830) 252. — L. auranthiacus (Waldst. and Kit.) Rchb. ibid. 
252, p. max. p. — Apargia aurantica Wald. and Kit. in Willd. Sp. pi. Ill, 
3 (1803) 1547.— A. crocea (Haenke) PoUini, Fl. Veron. II (1820) 565.— 
Ic: PoUini Ibid. t. 2, fig. 8; Hegi, 111. Fl. VI, 2 (1928) 1023, 1024; Javorka 
and Csapody, Icon. Fl. Hung. I (1929) 554, No. 3983. 

Perennial. Rhizome somewhat truncate, somewhat thick, with 
numerous fibrous roots; Stem-scape 15-30 cm high, with one, less often 
two capitula, with one to three (five) small linear leaves above, sulcate, 
glabrous, smooth below, with dense, simple, somewhat dark, hairs above 
mixed with white arachnoid pubescence, thickened below capitulum. Basal 
leaves three to six, 10-24 cm long, 0.5-2 cm wide, with simple, light- 
colored scattered hairs above, along margin and beneath along midrib, 
linear-lanceolate, acute, with straight or uncinate teeth. Capitula erect 
before and at flowering, 2-2.5 cm long, up to 3 cm wide. Involucral 
bracts imbricate, outermost bracts short, narrowly linear, middle ones 
linear-lanceolate, innermost bracts broadly linear; outer and middle bracts 
on back side, especially along midrib, covered with long, simple, dirty 
white or blackish, hairs; inner bracts glabrous, sometimes white-arachnoid 
along margin, often outside; corolla saffron-golden, with five concolorous 
teeth at apex; receptacle glabrous, flat. Achenes without beak, fusiform, 
ribbed, glabrous, slightly convex on one side, brown, 5 mm long, about 
1 mm wide, pappus as long as achene or slightly longer, dirty white, 
215 one-rowed, its bristles plumose above, thickened and barbed at base. 
Flowering second half of June to July; fruiting second half of July. (Plate 
XII, Fig. 1.) 

Patches of meadow and rocky outcrops in alpine zone. — European 
Part: Crimea(?), Upper Dniester. General distribution: central Europe, 
Mediterranean Region (Italy). Described from styrian Alps. Type in 
Vienna. 

Subgenus 2. Leontodon. — Apargia Scop. Fl. Camiol. ed. 2, II (1772) 
113, p. p.—Asterothrix Cass Diet. sc. nat. XL VIII (1828) 434.— Subgen. 
Apargia (scop.) Widder in Oesterr. Bot. Zeitschr. (1931) 146. Hairs forked 
or stellate or mixture of simple and forked hairs. 

Type of subgenus: type of genus. 



215 

Series 4. Asperi V. Vassil. — Root long, radish-shaped, only with 
forked and stellate hairs; achenes with long beak; involucral bracts with 
very short, white, somewhat thick, forked hairs along margin. 

10. L. asperus (Waldst. and Kit.) Boiss. Fl. or. Ill (1875) 730; Poir, 
Encycl. meth. Suppl. 453; Rchb. Fl. Germ. exc. 252. — Apargia aspera 
Waldst.and Kit. Icon. pi. Hung. II (1805) 114. — Leontodon crispus Vill. 
Hist. plfDauph. 3 (1789) 84.— L. biscutellaefolius DC. Prodr.VII (1838) 
103.— Ic: Rchb. Ic: Fl. Germ. XIX (1860) t.l370, II, 7-11; Waldst. 
and Kit op. cit. t. 110; Vill. op. cit. t. 25; Javorka and Csapody, Ic. Fl. 
Hung. I (1929) 554, No. 3977. 

Perennial. Root radish-shaped, merging with many-headed rhizome 
above. Stems 10-40 cm high, one-headed, less often with one or two 
branches and then with two or three capitula, scabrous from numerous 
short stiff setose, two- or three- (four)-forked hairs, with two to four 
small lanceolate-linear leaves, densely covered with erect forked hairs. 
Basal leaves narrowly lanceolate, narrowed toward base into winged 
petiole, sinuate-lobate, remotely toothed, hispid-scabrous on both sides 
from two to four-forked, very short hairs. Capitula usually turbinate before 
flowering, 1.5 cm long, up to 2 cm wide, up to 1.5 cm at fruiting, 
Involucral bracts, except innermost, covered with short, stiff, erect hairs 
along margin, with three- or four-forked hairs above, along keel, and 
with longer, setose, appressed hairs throughout on dorsal surface; inner 
bracts scarious along margin, entirely glabrous or with very fine appressed 
stellate hairs, as also middle involucral bracts. Receptacle covered with 
numerous white fimbriate scales; corolla light yellow. Achenes 8-12 mm 
long, flat on one side, weakly convex on the other, narrowed into a long 
216 beak above, sulcate, transversely rugose, setose- scabrous along beak; 
pappus two-rowed, up to 1 cm long, dirty white; bristles of outer row 
less numerous, barbed, shorter than inner; inner bristles plumose, not 
thickened at base, as also outer row of bristles Flowering second half of 
May to first half of August; fruiting June to August. (Plate XIII, Fig. 6.) 

Along dry stony and rocky slopes, meadows. — European Part: 
Volga-Kama (Kazan), Upper Dnieper, Middle Dnieper Volga-Don, 
Bessarabia, Black Sea Region, Crimea, Lower Don; Caucasus: 
?Ciscaucasia, Western and Southern Transcaucasia, Talysh. General 
distribution: Central Europe, Mediterranean Region, Atlantic Europe, 
Balkans-Asia Minor, Armenia and Kurdistan. Described from Hungary. 
Type in Budapest. 

Note. Intermediate forms between L. asperrimus and L. asperus are 
found in the Caucasus. Their origin can be established only by observing 
them in the field or after conducting crossing experiments with the 
hypothesized parent species. 



216 

ILL. asperrimus (Willd.) Boiss. Fl. or. Ill (1875) 12>2.—Scorzonera 
asperrima Willd. Sp. pi. Ill, 3 (1803) \5Q1 .—Apargia strigosa M.B. Fl. 
taur.-cauc. II (1808) 249. — Asterothrix asperrima (Willd.) Cass. Diet. sc. 
nat. XL VIII (1827) 434. — Podospermum asperrimum (Willd.) Sweet, Hort. 
Bot. ed. I (1827) 206. 

Perennial. Root radish-shaped, merging in many-headed rhizome 
above, bearing one to four (five) stems. Scapes 15-45 cm high, with a 
small number of small, linear or subulate, entire, acute leaves and with 
one fertile capitulum at apex. Sometimes one or two sparsely leafy twigs 
with apical capitulum, almost as long as stem, originate from lower half 
of main stem; stem base with dense rosette of oblong-lanceolate, entire 
or remotely sinuate-toothed leaves, 5-20 cm long, 0.5-2 cm wide; whole 
plant densely covered with long, stiff, straight, pale-green hairs, stellate 
above (with three or four upward-directed cusps). Capitula large, 2-2.5(3) 
cm long, 1.5 cm wide in middle and 2 cm wide above. Involucre 
cylindrical, bracts imbricate, Unear or lanceolate; outer bracts along keel 
and margin covered with stiff hairs, stellate-forked above; inner bracts 
flat, with membranous margin, as also others, weakly white arachnoid on 
dorsal surface. Receptacle weakly convex, densely covered with fimbriate 
scales. Achenes long, 15-20 mm long, fusiform, sulcate, somewhat 
flattened, transversely rugose, attenuate at apex into scabrous beak as 
long as achene; pappus dirty- white, of long (up to 1.5 cm), plumose bristles 
in two rows, and not thickened near base. Flowering first half of May to 
June; fruiting first half of June to July. (Plate XIII, Fig. 4.) 

Exposed stony and rocky mountain slopes, ravines. — Caucasus: 

Ciscaucasia, Eastern, Western and Southern Transcaucasia, Dagestan, 

Talysh; Soviet Central Asia: mountainous Turkmenia. General distribution: 

217 Asia Minor, Armenia and Kurdistan, Iran. Described from Galata. Type 

in Berhn. 

12. L. kotschyi Boiss. Diagn. pi. or. ser. I, 11 (1849) 40.— Ic: Hegi, 
111. Fl. VI, 2, t. 1026, 1030, 1031; Javorka and Csapody, Iconogr. Fl. 
Hung. t. 554. 

Perennial. Root long, radish-shaped, merging with one-, two- or three- 
headed rhizome above. Stems 7-30 cm high, simple, or, less often, with 
one or two branches and one to three capitula at apices of main stem and 
branches, with few linear or subulate-linear leaves, not thickened below 
capitulum, scabrous from very short, stellate, straight hairs. Basal leaves 
7.5-10 cm long, 0.8-2 cm wide, narrowly lanceolate, with winged petiole, 
sinuately pinnatifid or remotely toothed, with acute lobes and teeth, rather 
densely covered with very short, stiff, erect three- or four-forked hairs on 
both sides and along margin. Capitula obconical, 1.3-2 cm long, 1.12-2 
cm wide at flowering; involucral bracts imbricate, outer very short. 



217 

deflexed, lanceolate; inner bracts dense, appressed, linear-lanceolate, with 
very short stellate hairs along margin and midrib, remaining part finely 
arachnoid-hairy; inner involucral bracts scarious along margin, finely 
arachnoid-hairy on dorsal surface. Ligules yellow, unicolorous, with five 
long teeth; corolla tube and lower part of ligule sparsely sericeous. 
Receptacle convex, squamose, glabrous. Achenes up to 1 cm long, 
narrowed into beak above, sulcate, transversely undulate, sharply scabrous 
on beak, weakly flattened, brown; pappus dirty-white, as long as achene, 
two-rowed; bristles of outer row short, barbed, of inner row plumose, not 
thickened near base. Flowering June to July; fruiting July to August. (Plate 
XIII, Fig. 7.) 

Dry slopes, stony screes. — Caucasus: Talysh. General distribution: 
Iran. Described from Mt. Demavend [Damavand]. Type in Geneva. 

Series 5. Hispidi V. Vassil. Rhizome short, truncate; pubescence either 
exclusively of forked hairs (L. hispidum) or mixed, including simple and 
forked hairs; achenes without beak or with very short beak; involucral 
bracts arachnoid-hairy along margin in upper half. 

13. L. hispidus L. Sp. pi. (1753) 1124; DC. Prodr. VII, 102.— 
Hedypnois hispida (L.) Smith, Fl. Brit. 2 (1824) 813. — Apargia hispida 
(L.) Host, Fl austr. II (1831) 403.— Ic: Smith, op. cit. t. 554; Rchb. Ic. 
Fl. Germ. XIX (1860) t. 1368, 1369; Javorka and Csapody, Iconogr. Fl. 
Hung. I (1929) 554, No. 3980. 
218 Perennial. Rhizome short, somewhat truncate, horizontal or almost 

vertical, with numerous fibrous roots; scape straight, 10-60 cm high, 
scabrous from fine forked hairs, sometimes pinkish below, weakly 
thickened below capitulum, densely hairy, leafless or, less often, with 
one long leaf, mainly in upper part. Basal leaves 5-20 cm long, 1-3 cm 
wide, oblong or lanceolate, sinuate-toothed, incised-sinuate, covered 
with short, stiff, forked hairs, densely punctate-alveolate beneath with 
forked hairs on bottom, undulate-crisped in larger part, gradually narrowed 
toward base into rather long, sometimes pink, petiole. Capitula erect at 
flowering, drooping earlier, 1.2-3 cm wide at flowering; Ugules yellow. 
Involucral bracts linear, linear-lanceolate, dark green; outer bracts 
divergent, setose, bristles simple, mixed with some forked hairs, arachnoid- 
hairy along margin above; receptacle squamose, hairy, slightly convex. 
Achenes 4-6 mm long, 0.75 mm wide, fusiform, with relatively short 
beak, erect or slightly curved, longitudinally sulcate; transversely rugose, 
light brown, lustrous; pappus two-rowed; bristles of outer row short, 
barbed, inner bristles plumose, up to 7 mm long, thickened, lustrous at 
base. Flowering June to August; fruiting July to September. (Plate XIII, 
Fig. 2.) 



218 

Valley meadows, mountain slopes, hills. — European Part: Karelia- 
Lapland, Dvina-Pechora, Ladoga-Ilmen, Upper Volga, Volga-Kama, Upper 
Dnieper, Middle Dnieper, Volga-Don, Upper Dniester, Bessarabia, Black 
Sea Region, ?Crimea, Lower Don; Caucasus: Ciscaucasia, Dagestan, 
Eastern, Western and Southern Transcaucasia. General distribution: 
Scandinavia, Central Europe, Atlantic Europe, Balkans-Asia Minor. 
Described from western Europe. Type in London. 

14. L. caucasicus (M.B.) Fisch. Catal. Hort. Gorenk. (1812) 34; DC. 
Prodr. VII (1838) 104; Ldb. Fl. Ross. II, 7Sl. —Apargia caucasica M.B. 
Fl. taur.-cauc. II (1808) 247. 

Perennial. Rhizome reduced, with strong fibrous roots, very rarely 
creeping, long. Stem 14-50 cm high, glabrous, smooth, sulcate, slightly 
thickened and hairy only below capitulum, with one capitulum much 
longer than basal leaves and with 1-2 petiolate leaves. Basal leaves 3-20 
cm long, 1-2.5(3) cm wide, sinuate-toothed, runcinate, with triangular 
uncinate lobes, smooth above, glabrous or with simple hairs, sometimes 
mixed with forked hairs, glabrous beneath, or only along midrib covered 
with simple scattered long hairs mixed with forked hairs. Involucre dark 
green, with whitish or yellowish simple hairs, as also upper part of stem; 
outer bracts subulate, others lanceolate-linear, acuminate, sparsely hairy. 
Corolla bright yellow. Achenes dark brown, sulcate, more or less flattened, 
219 slightly curved, transversely rugose, toothed; pappus dirty white, sessile, 
two-rowed; bristles of outer row short, barbed; inner bristles plumose, 
longer than achene, weakly thickened toward base. Flowering July to 
August; fruiting August to September. 

Subalpine and alpine meadows, less often in forest zone in glades. — r 
Caucasus: Ciscaucasia, Dagestan, Eastern, Western and Southern 
Transcaucasia. Described from Caucasus. Type in Leningrad. 

Note. Undoubtedly close to L. hispidus L., from which it differs by 
having a smooth stem and smooth leaves with scattered simple hairs mixed 
with forked hairs. Apparently, a mountain form of L. hispidus L. 

GENUS1642. PicrisL.'^ 

L. Sp. pi. (1753) 792. 

Capitula many-flowered. Involucre imbricate, of lanceolate or linear 
bracts; outer bracts in several rows, short and recurved, inner bracts 
appressed, one- or two-rowed; outer bracts much shorter than inner. 



' Treatment by V.N. Vassilev. 

^ Derived from the Greek word picros — bitter. 



219 

Achenes weakly terete, thinner at both ends, longitudinally and transversely 
sulcate; beak absent or very short, almost one-third as long as achene, 
less often almost as long; pappus two-rowed, with outer row of shorter, 
almost filiform bristles, inner bristles plumose, connate at base. Herbaceous 
annuals, biennials, less often perennials, with erect branched stems, covered 
with stiff uncinate hairs. Leaves entire, toothed, less often pinnatisect. 

Type of genus: Picris hieracioides L. 

The genus includes up to 40 species, distributed in the Mediterranean 
Region, Eurasia, and northern Africa. 

1. Stem covered with simple as well as forked bristles or bristles 
with two anchor-shaped hooks above 2. 

+ Stem covered with simple as well as forked bristles with three or 
(often) four anchor-shaped hooks 6. 

2. Hairs light colored; corolla hmb two to three times as long as 
tube; achenes reddish-brown, slightly narrowed at both ends, very 
weakly falcate, without beak, oblong-sulcate, transversely undulate- 
rugose, 3 nmi long 1. P. hieracioides L. 

-I- Combination of characters different 3. 

220 3. Hairs black, dark green; corolla limb less than two times as long 
as tube; achenes dark brown, weakly falcate, narrowed at both 
ends, longitudinally sulcate, transversely cristate-rugose, 4 mm long 
2. P. japonica Thunb. 

+ Combination of characters different 4. 

4. Hairs black, somewhat dark but not light-colored; corolla limb 
slightly longer than tube; achenes weakly falcate, narrowed at both 
ends, with up to 5.5 mm-long weakly developed beak 

3. P. kamtschatica Ldb. 

+ Combination of characters different 5. 

5. Branches upward-spreading; inflorescence compressed, umbellate; 
stem roundish; hairs light colored, strongly thickened below; 
achenes light- or dark-brown, slightly falcate, narrowed at both 
ends, longitudinally sulcate, transversely undulate-rugose, 4 nmi 
long 4. P. rigida Ldb. 

+ Branches strongly divaricate, inflorescence lax; stem sulcate, hairs 
light colored, slightly thickened below; achenes brown, up to 5 
mm long, fusiform, with very short (0.3 mm long) beak, 

longitudinally sulcate, transversely undulate-cristate 

5. P. similis Vassil. 

6. Hairs with two anchor-shaped hooks; peduncles thickened above, 
abruptly narrowed directly below capitulum 

6. P. pauciflora Willd. 

+ Hairs with four hooks; peduncles equally thick throughout 7. 



220 

7. Plants strongly hairy; corolla limb as long as tube; achenes 5 mm 
long, fusiform, slightly falcate, longitudinally sulcate, dark, rather 

abruptly narrowed in up to 1.5 mm-long beak 

7. P. strigosa M.B. 

+ Plants sparsely hairy, mainly in lower part of stem; corolla limb 
two times as long as tube; achenes fusiform, 4 mm long, its beak 
up to 1 mm long, longitudinally sulcate, transversely undulate, 

with cristate membranous processes at apex and along beak 

8. P. canescens (Stev.) V, Vassil. 

Section Picris. — Sect. Eupicris Boiss. Fl. or III (1875) 734. Pappus 
of all achenes (outer and inner) similar. 
Type of section: type of genus. 

Series 1. Hieracioides V. Vassil. Biennial. Pubescence of simple 
forked hairs and hairs with two anchor-shaped hooks. Achenes without 
beak or with very short (0.2 mm long) beak. Peduncles uniformly thick 
throughout. 

221 1. P. hieracioides L. Sp. pi. (1753) 792; DC. Prodr. VII, 128; Ldb. 

Fl. Ross. II, 798; Kryl. Fl. Zap. Sib. XI (1949) 2910.— Leontodon 
umbellatus Schrank, Baier. Fl. II (1789) 334. — Apargia umbellata 
(Schrank) Schrank in Denkschr. Regensb. Bot. Gesellsch. II (1918) 66. — 
A. hieracioides (L.) Willd. Sp. pi. Ill, 3 (1803) 1554.— Picm aspera 
Gilib. Fl. lithuan. Ill (1785) 221 .—Hedypnois hieracioides (L.) Huds. Fl. 
Angl. (1762) 342.— Ic: Fedtsch. and Her. R. Evrop. Ross. (1910) 1041; 
Javorka and Csapody, Ic. H. Hung. I (1929) 555, No. 3987. 

Biennial. Stem erect, often more or less branched above, 50-100 cm 
high, as also leaves, covered with one-third to three-fourths mm-long, 
erect, stiff hairs, usually uncinate at apex. Basal leaves withering before 
anthesis; lower cauline leaves short-petiolate, linear-elliptical or broadly 
lanceolate, almost entire or sinuate-toothed, sometimes with narrow 
elongated teeth, 8-20 cm long, 0.5-6 cm wide; middle and upper leaves 
lanceolate, less often elliptical-lanceolate, sessile, semiamplexicaul, finely 
toothed, less often sinuate-toothed; uppermost leaves usually entire. 
Capitula in corymbose or corymbose-paniculate inflorescence, up to 1.5 
cm long and 1.5 cm wide at flowering and fruiting; involucral bracts 
dorsally covered with appressed arachnoid hairs and stiff, up to one-third 
to three-fourths mm long, appressed setose hairs, ciliate along margin. 
Florets bright yellow; corolla finely pubescent at base of limb; limb two 
to three times as long as tube; style pilose. Achenes yellowish or light 
brown, 3 mm long, falcate. Flowering July to August; fruiting August to 
September (Plate XIV, Fig. 2). 



221 

Forest, floodplain and steppe meadows, smooth stony slopes, forest 
edges, sometimes in crops. — European Part: Ladoga-Ilmen, Baltic Region, 
Upper Volga, Volga-Kama, Upper Dnieper, Upper Dniester, Middle 
Dnieper (northern part), Volga-Don, Trans- Volga Region (northern part); 
Caucasus: Ciscaucasia, Dagestan, Western and Southern Transcaucasia; 
Western Siberia: Ob', Upper Tobol, Irtysh; Soviet Central Asia: Aralo- 
Caspian, Balkhash Region. General distribution: Scandinavia, central and 
Atlantic Europe, Mediterranean Region, Balkans-Asia Minor, Armenia 
and Kurdistan, Iran. Described from western Europe. Type in London. 

Note. M.G. Popov (Ocherk Rast. i. Fl. Karpat. (1949) 254) segregated 
the eastern Carpathian specimens of this species as a separate subspecies: 
P. hieracioides ssp. carpatica M. Pop. 

2. P. japonica Thunb. Fl. Jap. (1784) 299; DC. Prodr. VII, 130; 
Ldb. Fl. Ross. II, 800, p. p., excl. pi. Kamtsch.; Kom. Fl. Manch. Ill, 2, 
766. — P. dahurica Fisch. ex Homem. Hort. Hafn. Suppl. (1819) 155; 
DC. Prodr. VII, 129. — Ic: Miyoshi and Makino, Pocket-Atlas Alp. pi. 
jap. II (1907) t. 66, fig. 381; Somoku Dzusetsu, ed. Makino (1912) t. 6. 

Biennial. Stem erect, usually slender, up to 1 m high, branched above, 
with sparse pubescence of stiff, erect, black hairs, majority with anchor- 
shaped tips. Cauline leaves sessile, oblong or oblong-lanceolate, acute, 
222 usually indistinctly toothed, less often sinuate-toothed, scabrous throughout 
from stiff setose hairs; basal leaves withering before anthesis. Peduncles 
slender; capitula up to 1.5 cm long, 1-1.5 cm wide, in lax corymbose 
panicle; involucral bracts dark green; peduncles and stem in upper part 
with fine appressed, soft, white, crisped hairs and also a large or small 
number of light colored or dark gray setose hairs, many with anchor- 
shaped tips. Corolla bright yellow; ligules less than two times as long as 
tube, with darker teeth; limb at base sericeous. Achenes 3-4 mm long, 
dark brown, longitudinally and transversely sulcate, slightly falcate, 
narrowed at both ends; pappus two-rowed, dirty white, readily detached; 
bristles of outer row short, barbed, forked; inner bristles longer, plumose. 
Flowering June to August; fruiting August to September (Plate XIV, 
Fig. 4). 

Dry meadows, meadow slopes, shrubby thickets. — Western Siberia: 
Ob', Irtysh, Altai; Eastern Siberia: Yenisei, Angara-Sayans, Dauria; Far 
East: Zeya-Bureya, Uda River area, Ussuri, Sakhalin, Kuril Islands; Soviet 
Central Asia: Balkhash Region, Dzhungaria-Tarbagatai, Tien Shan 
(northern). General distribution: China, Japan. Described from Japan. Type 
in Uppsala. 

Note. The ranges of P. japonica Thunb. and P. hieracioides L. are 
contiguous in western Siberia, Kazakhstan, and the Yenisei territory. As 
a result, plants with intermediate characters are found where they come 



222 

in contact. A form with long thin leaves and longer (up to 4.5 mm) 
achenes occurs in China (Hansu Province): var. chinensis V. Vassil. (foliis 
tenuioribus longioribusque acheniis longioribus). 

V.N. Woroshilov {Byull Glavn. Bot. Sada Akad. Nauk SSSR, 49, 
1963, 58) determined the distribution of Picris coreana (Kitam.) Worosch. 
{P. japonica var. coreana Kitam.) in the south of the Ussuri District. It 
also occurs in Korea and northeastern China. It differs from, P. japonica 
Thunb. by having a green involucre and light yellowish corolla. 

3. P. kamtschatica Ldb. in Mem. Acad. Sc. Petersb. V (1815) 557. 
— P. hieracioides subsp. kamtschatica (Ldb.) Hult. Fl. of Kamtch. IV 
(1930) 217.— P. japonica auct. non Thunb. Kom. Fl. Kamch. Ill (1930) 
185.— Ic: Takeda, Ic. Jap. alp. pi. t. 19. 

Biennial. Stem erect, up to 1 m high, thick, branched above, as also 
leaves, densely covered with erect, stiff, setose, blackish hairs, mostly 
with anchor-shaped tips. Basal leaves withering early; cauline leaves 
sessile, oblong, acute, indistincdy toothed. Capitula 2-2.5 cm long, up to 
2 cm wide, on densely pubescent, distinctly thickened peduncles above, 
in lax panicle or in compressed leafy corymb. Involucral bracts dark, 
223 two-rowed, pubescent with long erect blackish hairs; outer bracts short, 
usually divergent; inner bracts long, appressed. Corolla bright yellow; 
limb slightly longer than tube. Achenes 6-7 nmi long, scabrous, narrowed 
above into short beak; pappus dirty white, two-rowed, as long as achene, 
readily detached; brisdes of outer row shorter, with few branches; inner 
bristles longer, plumose. Flowering July to September; fruiting August to 
September. (Plate XIV, Fig. 3.) 

Coastal slopes and banks, landslides on mountain slopes; scattered.^ 
Far East: Kamchatka (and Bering Island). General distribution: Japan. 
Described from Kamchatka. Type in Leningrad. 

4. P. rigida Ldb. in Spreng. Syst. veg. Ill (1826) 663; Ldb. Fl. Ross. 
II 800. — P. hieracioides var. canescens Zing. Shorn. Sved. po Fl. Sredn. 
Ross. (1885) 271. 

Biennial. Stem erect, branched usually above, 30-80 cm high, as 
also leaves, especially in lower half, covered with stiff, simple or anchor- 
shaped white hairs with two hooks, at right angle to main stem and 
strongly thickened below. Leaves sinuate-toothed to entire; rosette of basal 
leaves usually persistent even after flowering; basal leaves lanceolate, 
linear-lanceolate, on winged petioles more or less as long as lamina; 
cauline leaves sessile, semiamplexicaul, lower ones broadly lanceolate, 
middle lanceolate, upper leaves linear-lanceolate, small. Capitula in 
terminal umbellate panicle, sometimes on very short peduncles (var. 
brachypoda DC), sometimes on axillary branches, up to 1.5 cm long at 



223 

1 cm wide, up to 12 cm long and 1.5 cm wide, at fruiting. Involucral 
bracts covered with short grayish fluff and light-colored, short, strongly 
thickened below, usually appressed setose hairs along margin and keel, 
imbricate; inner bracts narrowly lanceolate, appressed; outer bracts linear, 
sometimes strongly deflexed. Florets yellow; corolla tube at least one- 
third as long as limb; limb sericeous below; ligules with five narrow, up 
to 1 mm long, teeth. Achenes 4-5 mm long, longitudinally and transversely 
striate, slightly curved, narrowed at both ends, dark- or light-brown; pappus 
two-rowed; its brisdes connate at base, 7 mm long; outer bristles short, 
barbed; inner bristles long, plumose. Flowering July to September; fruiting 
August to September (Plate XIV, Fig. 1). 

Sands, gravels, stony and rubbly slopes, weedy places. — European 
Part: Upper Dnieper, Middle Dnieper, Volga-Don (southern part), Trans- 
Volga Region (southern part), Bessarabia, Black Sea Region, Crimea, 
Lower Don, Lower Volga, Western Siberia: Upper Tobol (southern part), 
Aralo-Caspian. Described from Crimea and Taganrog. Type and cotype 
in Leningrad. 

5. P. similis V. Vassil. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk 
SSSR XVII (1955) 459. 
224 Annual or biennial. Stem 10-130 cm high, solitary (or several arising 

from rootstock), usually divaricately branched, with long slender, more 
or less sulcate branches, with scabrous erect hairs having two anchor- 
shaped hooks. Leaves lanceolate, linear-lanceolate, linear, with stiff anchor- 
shaped and forked hairs; basal leaves persistent even after anthesis, 
narrowed into winged petiole; cauline leaves sessile, reduced and narrowed 
toward stem apex. Peduncles slender, uniformly thick throughout. Capitula 
in very lax panicles, small, oblong, 7-10 mm wide before flowering, up 
to 15 mm at fruiting, 1-1.5 cm long. Outer involucral bracts small, narrow, 
horizontal or deflexed; bracts of two inner rows as long as pappus, broadly 
scarious along margin, pubescent along keel; outer bracts arachnoid-hairy 
and with stiff hairs throughout. Achenes up to 5 mm long, about 1 mm 
thick, fusiform, longitudinally sulcate, transversely undulate-rugose, brown; 
pappus up to 6 mm long, two-rowed, of plumose hairs, connate below 
into ring. Flowering June to August; fruiting July to August (Plate XIV, 
Fig. 5). 

Stony, rubbly slopes, sands and gravelly areas in river valleys. — 
Soviet Central Asia: Aralo-Caspian, Balkhash Region, mountainous 
Turkmenia, Amu-Darya, Syr-Darya, Pamiro-Alai, Tien Shan. General 
distribution: China (eastern Tien Shan). Described from Soviet Central 
Asia. Type in Leningrad. 

Series 2. Pauciflorae V. Vassil. Annuals. Hairs simple and with two 
anchor-shaped hooks. Peduncles thickened upward, abrupdy narrowed 



224 

below capitulum. Achenes abruptly narrowed into up to 1 mm-long 
beak. 

Besides our species, this series includes P. kotschyi Boiss. from Asia 
Minor. 

6. P. pauciflora Willd. Sp. pi. Ill, 3 (1803) 1557. — Crepis lappacea 
Willd. op. cit. 1599. — Medicusia lappacea (Willd.) Rchb. Fl. Germ. exc. 
(1830) 254.— Ic: Rchb. Ic. Fl. Germ. XIX (1858) t. 1373. 

Annual. Stem 10-50 cm high, branched, scabrous from simple and 
anchor-shaped hairs with two hooks. Leaves 0.5-1 cm wide, oblong- 
lanceolate, toothed, sinuate-toothed or entire, with simple, forked and 
anchor-shaped stiff hairs; lower leaves petiolate, upper leaves sessile, 
lanceolate-linear. Peduncles thickened above, abruptly narrowed below 
226 capitulum. Capitula solitary at apices of branches 0.6-1 cm thick, 1-1.3 
cm long, somewhat constricted in middle at fruiting. Involucre imbricate; 
inner bracts compactly appressed, hnear-lanceolate, setose-ciliate, deeply 
furrowed at fruiting, with convex keel on back side; outer bracts linear, 
usually more or less deflexed or uncate. Corolla almost two times as 
long as involucre, pale yellow; limb almost two times as long as tube, 
with few hairs at base. Achenes 4 mm long, arcuate, narrowed at base, 
abruptly narrowed upward into 1 mm-long beak, with five deep 
longitudinal furrows, sharply transversely rugose; pappus plumose, two- 
rowed, with bristles connate at base; outer bristles shorter. Flowering May 
to June; fruiting June to July. (Plate XIV, Fig. 7.) 

Dry stony habitats, often as weed. — European Part: Crimea; 
Caucasus: Ciscaucasia, ?Dagestan, Western and Southern Transcaucasia. 
General distribution: central Europe, Atlantic Europe, Mediterranean 
Region, Balkans-Asia Minor, ?Armenia and Kurdistan, Iran. Described 
from France, Type in Berhn. 

Series 3. Strigosae V. Vassil. Annuals and biennials. Hairs simple 
and with four anchor- shaped hooks. Peduncles uniformly thick throughout. 
Achenes rather abruptly narrowed into 1.5 mm long beak. 

Besides the two species of our flora, this series also includes P. 
sprengeliana (L.) Lam. 

7. P. strigosa M.B. Fl. taur.-cauc. II (1808) 250, III (1819) 532; 
Grossh. Fl. Kavk. IV (1934) 223.— L. glaucescens DC. Prodr. VII (1838) 
130. 

Stem 25-80 cm high, strongly (sometimes from base) branched, four- 
or five-angled, with branches canescent from short appressed hairs, 
scabrous (especially below) from stiff, erect, anchor-shaped hairs with 
three or four hooks. Lower leaves oblong-lanceolate, runcinate, up to 



225 




225 Plate XIV 

1 — picris rigida Ldb.; 2 — P. hieracioides L.; 3 — P. kamtschatica Ldb.; A — P. japonica 
Thunb.; 5 — P. similis V. Vassil.; 6 — P. strigosa M.B.; 1—P. pauciflora Willd. 



226 

10 cm long, petiolate; upper and middle leaves linear-lanceolate, at base 
narrowed, not amplexicaul; all leaves more or less covered with anchor- 
shaped and forked light-colored hairs. General inflorescence with many, 
scattered, small capitula at apices of branches, 5-6 mm wide before 
flowering, up to 1 .5 cm wide and about 1 cm long at anthesis and fruiting. 
Capitula many-flowered. Involucre, besides gray tomentum, covered with 
stiff, setose, simple and anchor-shaped, hooked hairs, two-rowed; outer 
bracts narrow, deflexed; inner bracts broader, appressed, scarious along 
margin. Florets pale yellow; limb at base pubescent, as long as corolla 
tube, ligules with five darker teeth. Achenes 5 mm long, longitudinally 
227 sulcate, transversely undulate, dentate above, fusiform, weakly arcuate, 
dark, rather abruptly narrowed into short (up to 1.5 mm-iong) beak; pappus 
dirty-white. Flowering May to August; fruiting June to September. (Plate 
XIV, Fig. 6.) 

Rocks, dry slopes and meadows. — European Part: Crimea; Caucasus: 
Ciscaucasia, Dagestan, Eastern, Western and Southern Transcaucasia, 
Talysh; Soviet Central Asia: mountainous Turkmenia (Kopetdag). General 
distribution: Balkans-Asia Minor, Armenia and Kurdistan, Iran. Described 
from the Caucasus. Type in Leningrad. 

8. P. canescens (Stev.) V. Vassil. comb. nova. — P. laevis C.A. Mey. 
Verzeichn. Pfl. Cauc. (1831) 59; Ldb. Fl. Ross. II, 3, SOI.— Hypochaeris 
canescens Stev. in Mem. Soc. Nat. Mosc. IV (1813) 100 and in M.B. Fl. 
taur.-cauc. IV (1819) 539. — Dickera canescens (Stev.) C.A. Mey. Ind. 
Sem. Hon. Petrop. IV (1837) 34. 

Annual or biennial. Stem erect or ascending, strongly branched, often 
from base, 25^0 cm high, with basal rosette of leaves, usually as also 
leaves, glabrous or only at base with stiff erect hairs, with two, three or 
four anchor-shaped hooks at tip. Leaves more or less linear, sinuate- 
toothed; upper leaves linear, entire; basal leaves petiolate; cauline leaves 
short-petiolate or sessile, 3-10 cm long, 0.3-2.5 cm wide. Capitula in 
panicles, numerous, up to 2 cm long and up to 1.5 cm wide at flov/ering. 
Involucral bracts flat, with appressed arachnoid pubescence, less often 
with a few stiff, setose, simple hairs. Florets yellow; corolla pubescent at 
limb base; corolla tube half as long as limb. Style short-setose. Achenes 
up to 4 mm long, dark brown, weakly arcuate, fusiform, with short (up 
to 1 mm) beak, longitudinally sulcate, transversely undulate, with cristate 
projections at apex and along beak. Flowering July to August; fruiting 
July to September. 

Along riverbanks, in stony places. — Caucasus: Ciscaucasia (eastern), 
Dagestan. Described from Ciscaucasia (Terek Valley). Type in Leningrad. 

Note. It is close to P. strigosa M.B., with which it shares the presence 
of hairs with four anchor-shaped hooks, but well differentiated from it in 



227 

being weakly hairy throughout (sometimes completely lacking 
pubescence), and having flat involucral bracts, and a shorter achene with 
a shorter beak. The possibility of a hybrid origin (P. strigosa M.B. x, P. 
rigida Ldb.) cannot be excluded. 



GENUS 1643. Helminthia Juss' ^ 

Juss. Gen. pi. (1774) 170. 

Involucre two-rowed; inner row of eight narrow bracts; outer row of 
three to five small, wide bracts. Receptacle without scales. All florets 
228 ligulate, yellow. Achenes flattened-terete, transversely striate, obtuse in 
upper part, rugose, narrowed into filiform beak; pappus one-rowed, its 
bristles simple, persistent. Leaves oblong, semiamplexicaul. Stem strongly 
branched. Leaves and capitula with stiff setose hairs. Annuals or biennials. 

Type of genus: Helminthia echioides (L.) Gaertn. An oligotypic 
Mediterranean genus. 

1. H. echioides (L.) Gaertn. De fruct. II (1791) 368. — Picris echioides 
L. Sp. pi. (1753) 792. — Helminthia humifusa Trev. in Nova Acta Acad. 
Caes. Leopold. Carol. Nat. Cur. 13, I (1826) 195.— Ic: Gaertn. op. cit. t. 
159, f. 2; Javorka and Csapody, Ic. Fl. Hung. I (1929) 555, No. 3985. 

Annual. Stem 3-8 mm thick, erect or prostrate, often branched from 
base, covered with uncinate or forked stiff setose hairs. Leaves oblong- 
lanceolate, entire or sinuate-toothed; basal leaves petiolate; cauline leaves 
amplexicaul, auriculate. Capitula crowded in groups of two to five at 
branch apices, on short peduncles. Outer involucral bracts oval-cordate, 
acuminate, inner narrowly linear, subulately long-acuminate, stiff-hairy 
or ciliolate. Achenes oblong, finely rugose, abruptly narrowed above into 
slender, fragile beak, patelliform-broadened above, with white, one-rowed 
pappus of plumose hairs. Flowering June to July; fruiting July to August 
(Plate XII, Fig. 7). 

A weed in cultivated fields, roadsides. — European Part: Crimea; 
Caucasus: Dagestan, Eastern Transcaucasia; Soviet Central Asia: 
mountainous Turkmenia (introduced?). General distribution: central 
Europe, Atlantic Europe, Mediterranean, Balkans-Asia Minor, Armenia 
and Kurdistan, Iran and Afghanistan, North and South America 
(introduced?). Described from western Europe. Type in London. 



' Treatment by V.N. Vassilev. 

^ From the Greek word elmis — worm; so named on the basis of the striate achenes. 



228 

GENUS 1644. Urospermum Scop.' ^ 

Scop. Introd. hist. nat. (1777) 122. 

Capitula solitary, large, at apices of main stem and lateral branches. 
Involucral bracts similar, eight, connate at base. Receptacle glabrous, 
alveolate. All florets ligulate, grayish-yellow, with five brown teeth at 
apex; outer florets with purple tinge. Achenes transversely striate, curved, 
with hollow basally swollen beak; pappus one-rowed, its bristles plumose. 
Annual herbs; stem simple or with few lateral branches, glabrous or with 
stiff hairs, leafy; lower leaves petiolate, middle and upper leaves sessile, 
amplexicaul, toothed, pinnatifid or entire. 

Type of genus: U. picroides (L.) Desf. 
229 An oligotypic genus, comprising only four species, three of which 

are confined to the Mediterranean Region, and one is occurs in South 
Africa. 

1. U. picroides (L.) Desf. Catal. Hort. Paris, ed. 1 (1820) 90; Ldb. 
H. Ross. Ill 790; Boiss. FI. or. Ill, 743.— f/. asperum (L.) DC. Fl. France, 
ed. 3, IV, 1 (1805) 63.— Tragopogon picroides L. Sp. pi. (1753) 790.— 
T. asperum. L. Ibid. 790. — Arnogon picroides Willd. Sp. pi. Ill (1803) 
1496.— A. asper Willd. op. cit. 1497.— Ic: Rchb. Ic. Rchb. Ic. Fl. Germ 
XIX, t. 1377. 

Annual. Stem erect, 15-40 cm high, hispid; simple or weakly 
branched. Leaves thin; lower leaves petiolate, oblong-spatulate, lyrate; 
middle and upper leaves sessile, amplexicaul. auriculate, oblong or ovate, 
acutely sharp-toothed, pinnatifid, less often entire, hispid beneath along 
veins. Capitula (one) three to five, on long peduncles, at apices of main 
stem and branches, large, 1.7-3.0 cm long, up to 15 mm wide at flowering, 
up to 3 cm wide with mature fruits; involucre one-rowed, of eight basally 
connate, up to 2 cm long, lanceolate bracts with narrow white membranous 
margin, glabrous or with few setaceous hairs; ligules pale yellow, with 
five apical teeth; receptacle without scales, alveolate. Achenes with 
pappus up to 2.5 cm long; pappus readily detached, one-rowed, of white 
plumose bristles; beak, up to 11 mm long, swollen below, hollow, dark 
brown, making obtuse angle with achene; achenes without beak and 
pappus 4-5 mm long, curved, flat, with punctate tubercles along margin, 
light brown. Flowering April to June; fruiting May to June. (Plate XII, 
Fig. 5.) 

Mountain slopes on stony soils, among shrub thickets and citrus 
plantations. — Caucasus: Eastern Transcaucasia. General distribution: 



' Treatment by V.N. Vassilev. 

^Name derived from the Greek words ura — tail, and sperma — seed. 



229 

Mediterranean Region, Balkans-Asia Minor, Armenia and Kurdistan, Iran. 
Described from southern Europe. Type in London. 

GENUS 1645. Hedypnois Schreb.'-^ 

Schreb. Gen. pi. II (1791) 532. 

Capitula many-flowered, solitary. Receptacle glabrous. Involucre two- 
rowed; outer involucral bracts few, short; inner bracts several times as 
long as outer, at fruiting hardening and enclosing outer achenes. Ligulate 
florets yellow, exceeding involucre. Outer achenes slightly curved, terete, 
finely ribbed, with toothed membranous crown at apex; inner achenes 
straight, with pappus of lanceolate-subulate serrate bristles. 

230 Type of genus: H. cretica (L.) Willd. 

1. Involucral bracts glabrous, smooth 1. H. cretica (L.) Willd. 

-I- Involucral bracts hispid 2. H. persica M.B. 

1 H. cretica (L.) Willd. Sp. pi. Ill, 3 (1800) 617; DC. Prodr. VII, 1, 
81; Ldb. Fl. Ross. II, 2, 773; p. p. Boiss. Fl. or. Ill, 719; Grossh. Fl. 
Kavk. IV (1935) 225.— Hyoseris cretica L. Sp. pi. (1753) 810. 

Annual. Stems usually dichotomously branched, less often simple, 
with scattered hairs, sometimes almost glabrous. Basal and lower cauHne 
leaves oblong or oblong-spatulate, on winged petiole, hispid, more or 
less toothed, to sinuate-toothed; cauline leaves oblong-lanceolate, sessile. 
Capitula solitary, at apices of branches and main stem, on long, upwardly 
thickened peduncles, fistular above. Involucral bracts oblong-lanceolate, 
glabrous or finely scatteredly puberulent, hardening at fruiting, carinate, 
enclosing outer achenes. Achenes linear-cylindrical, 6-7 mm long; outer 
achenes with short, membranous, toothed corona at apex, inner achenes 
with pappus of membranous, lanceolate-subulate, finely toothed, 3-4 mm 
long bristles. Flowering April to May; fruiting May to June (Plate I, 
Fig. 1). 

Roadsides, in gardens, vineyards, on clayey slopes, sometimes in 
fields. — European Part: Crimea (southern coast); Caucasus: Eastern and 
Southern Transcaucasia, Talysh. General distribution: Mediterranean, Asia 
Minor. Described from Island of Crete. Type in London. 

Note. This species is more mesophilous than H. persica. In Crimea, 
it is found in the coastal belt in the south and in the Caucasus only under 
high moisture conditions (irrigated fields, etc.). 



' Treatment by I.T. Vassilczenko. 

^ From the Greek words hedys — sweet, and pnois — smell (on the basis of the aromatic 
odor of the plant). 



230 

2. H. persica M.B. Fl. taur.-cauc. Ill (1819) 539; Boiss. Fl. or. Ill, 
119.— H. cretica auct. non Willd.; Ldb. Fl. Ross. II, 2 (1845-1856) 733, 
p. p. 

Annual. Stem 5-25 cm high, leafy, usually branched from base; all 
plant parts (except corolla) hispid. Basal leaves oblong, on long and 
thickened petioles, sinuate-toothed, with triangular remote teeth sometimes 
weakly developed; cauline leaves oblong-lanceolate, amplexicaul, sessile, 
sinuate-toothed, but with narrower teeth. Capitula solitary, at apices of 
main stem and its branches, on long, upwardly clavately thickened and 
fistular peduncle. Involucral bracts densely hispid, carinate at fruiting, 
hardened, with whitish pappillae and spinules in middle on dorsal surface, 
densely covered with long stiff hairs and shorter hairs throughout on outer 
surface, inner side glabrous, enclosing outer achenes. Achenes linear- 
231 cylindrical, finely pilose, with indistinct longitudinal ribs, dark brown; 
outer achenes with erose-toothed corona; inner achenes with pappus of 
membranous bristles. Flowering May. 

Dry stony and clayey slopes, in plains and foothills, sometimes as 
weed of roadsides, among crops, etc. — Caucasus: Eastern and Southern 
Transcaucasia, Dagestan. General distribution: Iran, Balkans-Asia Minor, 
eastern Mediterranean Region. Described from Derbent. Type in 
Leningrad. 



GENUS 1646. Garhadiolus Jaub. and Sp 



1,2 



Jaub. and Sp. Ill, pi. or. Ill (1847-1850) 119. 

Capitula few-flowered, short-cylindrical. Receptacle glabrous; 
involucre two-rowed, of inner long, linear-lanceolate bracts and outer very 
small, inconspicuous bracts, latter hardening at fruiting, incurved along 
margin and more or less firmly enclosing lower part of outer achenes. 
Ligules yellowish, slightly lower than involucral bracts. Achenes terete, 
thickened below, more or less curved, with short pappus at apex. Annual 
herbs with simple, more or less toothed (to lobate and pinnatifid) leaves. 

Type of genus: G. angulosus Jaub. and sp. 

The genus includes four or five species, out of which two are found 
in the USSR. 

1 Inner achenes with long beak and somewhat long pappus; 
peripheral achenes narrowed at apex, with corona of very short 
bristles, bend of achene closer to its base; inner involucral bracts 



Treatment by I.T. Vassilczenko. 
■ Anagram of Rhagadiolus (cf.). 



231 

usually spinose, sometimes lacking spines 

1. G. papposus Boiss. and Buhse 

+ All achenes narrowed at apex and with small fimbriate corona; 
peripheral achenes bent (at right angle) above; involucral bracts 

smooth or, less often, more or less setose 

2. G. angulosus Jaub. and sp. 

1. G. papposus Boiss. and Buhse in Nouv. Mem. Soc. Nat. Mosc. 
XII (1860) 135; Boiss. Fl. or. Ill, 723; Grossh. Fl. Kavk. IV (1935) 226; 
Nevski in Som. Rast. SSSR IV, 318; O. and B. Fedtsch. Perech. Rast. 
Turk. 4 (1911) 305; Kovalevskaja in Fl. Uzb. VI (1962) 426.— G. 
scaberrimus Franch. ex O. and B. Fedtsch. Perech. Rast. Turk. 4 (1911) 
306.— G. echinatus Pavl. in Vestn. Akad. Nauk Kazakh. SSR 3 (1950) 
27. — Rhagadiolus papposus O. Ktze. in Tr. Peterb. Bot. Sada X (1887) 
205. — Koelpinia scaberrima Franch. in Ann. Sc. Nat. VI ser. 15, (1883) 
331. — Ic: Boiss. and Buhse op. cit. t. 7; Kovalevskaja op. cit. Plate 51, 
Fig. 1. 
232 Annual. Stems usually branched, ascending, 5-40 cm high, more or 

less hairy or almost glabrous, sometimes whole plant hispid (var. 
scaberrimus (Franch.) Vass.), Basal and lower cauline leaves oblong or 
oblong-ovate, petiolate, more or less unevenly sinuate-toothed to lyrately 
pinnatilobate and pinnatipartite, setose at least beneath along veins; cauline 
leaves like basal, but sessile and reduced. Capitula terminal and axillary. 
Inner involucral bracts lanceolate, more or less convexo-concave, usually 
setose on outer side (sometimes glabrous), and often also arachnoid- 
woolly, hardening at fruiting, carinate, short, more or less setose-spinose 
on dorsal side, tuberculate or smooth (var. suberculatus Vass.), enclosing 
outer achenes, latter more or less bent (to more or less straight), with 
slender beak and pappus of short, white, finely barbed bristles; all achenes 
puberulent, hairs appressed. Flowering March to April. 

Plains and foothills. — Caucasus: Ciscaucasia (eastern part), Dagestan, 
Eastern and Southern Transcaucasia; Soviet Central Asia: All regions. 
General distribution: Iran, Armenia and Kurdistan. Described from 
southern Transcaucasia. Type in Geneva. 

Note. The hispid plants have been treated by some authors as a 
separate species (C. scaberrimus Franch., G. echinatus Pavl.), which is 
not justified, as it is only the variety typical of more or less light soils 
(sand, stony soils, etc.). The Herbarium of the Botanical Institute, Academy 
of Sciences of the USSR, has plants which were identified as G. 
scaberrimus Franch. by B.A. Fedtschenko already in 1915. Besides this 
hispid variety, there is another variety with smooth outer involucral bracts, 
as if made of cork (from which the name var. suberculatus is derived). 
This variety is found under conditions of a better and, apparently, sure 



232 

moisture supply. Both varieties are found throughout the range of the 
species and are connected to each other and to typical G. papposus by 
intermediates. 

2. G. angulosus Jaub. and Sp. III. pi. or. Ill (1849) 12; Koyalevskaja 
in Fl. Uzb. VI (1962) 427.— G. hedypnois (Fisch. and Mey.) Jaub. and 
Sp. 111.' pi. or. Ill (1849) 120; Boiss Fl. or. Ill, 723; Grossh. Fl. Kavk. IV 
(1934) 225; Nevski in Som. Rast. SSSR IV, 318.— G. minutissimus 
Kitamura in Acta Phytotax. et Geobot. XVII (1957) 75 Rhagadiolus 
hedypnois Fisch. and Mey. ex DC. Prodr. VII (1938) 78 and in Ind. Sem. 
hort. Petrop. IV (1838) 46; Ldb. Fl. Ross. II, 2, 772; O. and B. Fedtsch. 
Perech. Rast. Turk. 4 (1911) 305, non All. (1785). Koelpinia hedypnois 
H. Baill. in Ann. Sc. Nat. ser. 16 (1883) 330. — Hedypnois minutissima 
Bge. in Mem. Ac. Sc. Petersb. VII (1851) 372. — Ic: Jaub. and Sp. op. 
cit. t. 284. 
233 Annual. Stem simple or dichotomously branched, 5-20 cm high, 

pubescent with almost scaly hairs below (or throughout). Basal and lower 
cauline leaves oblong-obovate, more or less sinuate-toothed to divided, 
petiolate; cauline leaves ovate or obovate-lanceolate, sessile, toothed; all 
leaves glabrous or hispid (mostly along margin and along veins beneath). 
Capitula small, terminal on peduncles thickened in upper part; axillary 
capitula almost sessile. Involucral bracts glabrous or densely pubescent 
(van lipskyanus m.); outer involucral bracts conduplicate at fruiting, 
hardened, 7-9(11) mm long, glabrous or setose. Achenes 4-6 nrni long, 
about 4 mm wide, grayish-dark-brown, indistinctly longitudinally ribbed, 
somewhat remote (narrow flabellate above in capitulum); peripheral 
achenes half to two-thirds enclosed in involucral bracts, bent above in 
free part almost at right angle; inner achenes free, thicker, curved (to 
annular); achenes more or less finely setose (inner or all) or glabrous, 
with fimbriate corona at apex. Flowering April to May. 

Loess plains, foothills (extending up to 1,500-2,000 m), usually as 
weed in fields. — Caucasus: Eastern and Southern Transcaucasia; Soviet 
Central Asia: Syr-Darya, Amu-Darya, Pamiro-Alai, Tien Shan, 
mountainous Turkmenia. General distribution: West Asia (including 
Afghanistan). Described from Transcaucasia, from the Beshbarmak 
Mountains. Type in Leningrad. 

GENUS 1647. Rhagadiolus Scop.'^ 
Scop. Introd. (1777) 122. 



' Treatment by I.T. Vassilczenko. 

- From the Greek word rhagas — slit (involucral bracts with slit-like, indistinct 
depression). 



233 

Capitula many-flowered, Receptacle glabrous. Involucre two-rowed; 
outer involucral bracts five, small, ovate-triangular, concavo-convex, inner 
bracts five to eight, much longer, linear, in fruit expanded and hardened, 
enclosing outer, stellately arranged achenes. Corolla yellow. Outer achenes 
linear-subulate, narrowed above, more or less falcate; inner achenes terete, 
usually strongly curved, up to annular; all achenes without pappus. 

Type of genus: R. edulis Gaertn. 

The genus includes five or six species distributed in the Mediterranean 
Region and West Asia. 

1 . Basal and lower cauline leaves lyrately pinnatipartite or pinnatisect; 

involucral bracts smooth 1. R. edulis Gaertn. 

234 + Basal and lower cauline leaves more or less sinuate-toothed, entire; 
involucral bracts dorsally setose ... 2. R. hebelaenus (DC.) Vass. 

1. R. edulis Gaertn. De Fruct. et Sem. PI. II (1791) 354; Willd. Sp. 
pi. Ill, 1625; M.B. Fl. taur.-cauc. Ill, 541. — R. lampsanaefolius Mirb. 
Hist. Nat. PI. XI (1805) 328.— R. stellatus T. edulis DC. Prodr. VII, 1 
(1838) 77; Ldb. Fl. Ross. II, 2, 772. — Lampsana rhagadiolus L. Sp. pi. I 
(1753) 912.— Exs.: Dorfler Herb. Norm. No. 3060 (sub R. rhagadiola 
(L.) Dorfl). 

Annual. Stem 5-30(40) cm high, simple or weakly branched, glabrous. 
Basal (and lower cauline) leaves with winged petioles, lyrately 
pinnatipartite or pinnatisect; termial lobe large, brownish, ovate-reniform, 
more or less angular, lateral lobes much smaller, broadly ovate; leaves 
scatteredly hairy beneath and along petiole; cauline leaves sessile or 
subsessile, oblong-rhombic or oblong-lanceolate, more or less sinuate- 
toothed; uppermost leaves narrow-oblong, weakly toothed to almost entire. 
Capitula small, apical and axillary; involucral bracts smooth; florets 
exceeding involucre by one-third to three-fourths. Achenes stellately 
arranged, linear-terete; outer achenes enclosed in hardened and connate 
involucral bracts, straight, usually weakly uncinate only at apex, 8-15 
mm long; inner achenes annular, densely and finely pubescent. Flowering 
April to May. 

A weed of roadsides on slopes, near fences, buildings, in kitchen 
gardens, etc.; apparendy only an adventive in our flora. — European Part: 
Crimea; Caucasus: Western Transcaucasia. General distribution: 
Mediterranean Region, Balkans-Asia Minor. Described from the Orient. 
Type in London. 

2. R. hebelaenus (DC.) Vass. comb. nov. — R. stellatus d. hebelaenus 
DC. Prodr. VII, 1 (1838) 78; Boiss. Fl. or. Ill (1875) 722. 

Annual. Plants 5-25(30) cm high with weakly branched stem, 
puberulent, more sparsely above, sometimes almost glabrous. Basal and 



234 

lower cauline leaves on wide (winged) petioles, more or less sinuate- 
toothed, to pinnatifid and divided, puberulent; upper cauline leaves sessile, 
lanceolate or oblong-lanceolate, toothed to entire, finely hairy. Capitula 
terminal and axillary. Inner involucral bracts dorsally covered with light- 
colored, erect scaly brisdes. Peripheral achenes terete, stellately arranged, 
(10)12-15 mm long, more or less falcate (resembling achenes of Koelpinia 
linearis in form, but smooth), enclosed in expanded and hardened 
involucral bracts with bristles or tubercles on dorsal surface; inner achenes 
terete, more or less annular, densely and finely pubescent. Flowering 
April to May. 
235 Cultivated fields, dry slopes, roadsides, plains and lower foothills 

(up to 200-300 m). — Caucasus:, Dagestan, Eastern and Southern 
Transcaucasia. General distribution: eastern Mediterranean, Balkans-Asia 
Minor. Described from Greece. Type in Paris. 

Subtribe 5. Lactucinae Less. Synops. Compos. (1832) 135 
("Lactuceae"); DC. Prodr. VII, 1 (1838) 133; Benth. in Benth. and Hook, 
f. Gen. pi. II (1873) 168, 222. — Trib. Lactuceae sect. 1. Archetypae Cass. 
in Diet. sc. nat. XXV (1822) 60, p. min. p.; sect. 2. Crepideae Cass. op. 
cit. 61. p. p.; sect. 3. Hieracieae Cass. op. cit. c. 63, p. p. — Trib. Lactucees 
sect. 1. Prototypes III. Prototypes vraies Cass, in Diet. sc. nat. XLVIII 
(1827) 422; sect. 2. Crepidees II. Crepidees vraies Cass. ibid. sect. 3. 
Hieraciees Cass. ibid. p. max. p. — Trib. Lactuceae subtrib. Lactuceae 
archetypael (Cass.) Dum. Fl. Belg. (1827) 59; subtrib. Crepideae (Cass.) 
Dum. ibid. p. max. p. — Subtrib. Hieracieae (Cass.) Dum. op. cit. 62. — 
Trib. Cichorieae. c) Hieracieae, d) Crepideae, e) Lapsaneae and f) 
Lactuceae Dum. Analyse fam. pi. (1829) 30. — Trib. Hieraceae D. Don 
in Edinb. New Philos. Joum. VI (1829) 306.— Trib. Lactuceae D. Don 
op. cit. 307. — Subtrib. Lampsaneae Less. Synops. Compos. (1832) 126, 
p. min. p.; DC. 1. c. (eodem ambitu). — Subtrib. Hieracieae Less. op. cit. 
140; DC. op. cit. 198.— Gruppe Chondrilleen W.D. Koch, Taschenb. 
Deutsch. u. Schweiz. Fl. (1844) 314. — Gruppe Lactuceen W.D. Koch op. 
cit. 316. — Gruppe Crepideen W.D. Koch op. cit. 318. — Subtrib. Lapsaneae 
Benth. in Benth. and Hook. f. Gen. pi. II (1873) 168, 220, p.p.— Subtrib. 
Rhagadioleae Benth. ibid. p. max. p. — Subtrib. Crepideae Benth. op. cit. 
168, 221, p. max. p. — Subtrib. Hieracieae Benth. Ibid. — Subtrib. 
Hypochoerideae Benth. op. cit. 168, 222, p. min. p. — Subtrib. Cichorinae 
O. Hoffm. in Engl. u. Prantl. Nat. Pflanzenfam. IV, 5 (1893) 353, p. min. 
p. — Subtrib. Crepidinae O. Hoffm. op. cit. 366; Stebbins in Madrono 
XII, 3 (1953) 76. — Pappus usually of numerous, thin, mostly white, smooth 
or weakly barbed hairs (under lens!), sometimes unequal in length, more 
rarely also in thickness; in other cases pappus of outer tuft of short reduced 
hairs (under lens!) and inner normal hairs; very rarely pappus weakly 



235 

developed or entirely absent. Pollen grains usually spiny-cristate 
(echinolophate). Receptacle glabrous, smooth or somewhat tuberculate, 
alveolate or punctate; less often alveolate with hairs, ciliolate or fimbriate. 
Basic chromosome number: x = 9, 8, 7, 6, 5, 4 and 3. 

Type of subtribe: genus Crepis L. 

Crepidinae is the largest subtribe of the Cichorieae. Several major 
lines are clearly distinguishable in its phylogenetic development. 

1. Launaea — Sonchus Line (from the genus Paramicrorhynchus to 
the genus Reichardia). This mainly includes the African genera and species 
with pappus hairs not always similar in length and thickness. Plants usually 
of dry habitats. 

236 2. Hieracium Line. Includes only two genera (Hieracium and 

Andryala). Achenes cyhndrical and truncate. Pappus hairs mostly fragile, 
usually not pure white. 

3. Prenanthes — Lactuca Line (from the genus Prenanthes to the 
genus Lagoseriopsis; however, the referral of the latter genus to this line 
is still not altogether justified). Pappus hairs numerous, more or less soft 
and white, rarely dirty white or rusty, denticulate (visible under hand 
lens!) to almost smooth, or pappus dimorphic: outer tuft of short reduced 
hairs (under hand lens!), inner normal hairs. The most primitive members 
of this line are mesophytic herbs, growing in forests in the subalpine 
zone of the temperate regions of Eurasia. Mostly quite leafy, often tall, 
herbaceous plants. 

4. Youngia — Ixeris Line (from the genus Youngia to the genus 
Acanthocephalus). Mostly smooth or weakly pubescent plants; achenes 
usually tuberculate, especially at the apex. Pappus usually of barbed hairs, 
sometimes reduced or entirely absent. 

5. Crepis Line (the genera Crepis, Lagoseris and Zacyntha). Plants 
usually pubescent, sometimes glabrous; hairs soft, sometimes stiff or 
setose. Pappus usually present (very rarely absent in marginal achenes), 
of scabrous-serrate or dentate, soft, generally white hairs. The central 
genus Crepis L. includes species distributed in the Old as well as the 
New World. 

GENUS 1648. Paramicrorhynchus Kirp. Gen. Nov.' ^ 

Kirp. in Addenda XXVIII, 725. 



' Treatment by M.E. Kirpicznikev. 

^ From the Greek word para — near, and the generic name Microrhynchus. 



236 

Capitula homogamous, with 15-30 florets, usually cylindrical, solitary 
in groups of few capitula at apices of branches and in axils. Involucre 
(three-) four or five-rowed, imbricate; bracts scarious along margin. 
Achenes heteromorphic: outer achenes brown or gray, with rather 
numerous, prominent, longitudinal, tuberculate-rugose ribs, narrowed at 
base, deltoidly narrowed at apex, terminating into slender, scarcely visible, 
fragile beak; inner achenes prismatic-columnar, yellowish, with few 
(usually five) deep longitudinal furrows, somewhat membranous-rugose 
or smooth; intermediates between these two main types also found in 
each capitulum. Pappus of numerous thin, silky, white, somewhat barbed 
(under lens!) hairs, connate into a ring. 

Type of genus: Paramicrorhynchus procumbens (Roxb.) Kirp. 
237 Note. This species, referred by me to the new genus 

Paramicrorhynchus, is referred by the majority of contemporary 
taxonomists to Launaea Cass, and by some to Microrhynchus Less. The 
type species of Launaea — L. bellidifolia Cass. — grows on coastal sands 
of Madagascar. This plant is characterized by runner-like shoots and 
homomorphic achenes with five very prominent, semi-terete ribs. Our 
species cannot be retained in the genus Launaea, even if only because 
the achenes of the latter are homomorphic. Much more complex is the 
problem of the relationship of the species in question to Microrhynchus 
Less. This genus was established with a single species, M. nudicaulis 
(L.) Less. The basionym of the combination was Chondrilla nudicaulis 
L. Linnaeus in his original description stated: "Habitat in American 
septentrionali; ad pyramides aegypti." There is every reason to believe 
that "North America" is indicated here by mistake (misprint?) instead of 
"North Africa." Causing much greater difficulty is the fact that the original 
description, the literature, and the herbarium material do not yet allow us 
to determine precisely which particular plant under the name C. nudicaulis 
corresponds to the authentic specimen of Linnaeus. An annual plant figures 
in the works of some (mainly older) authors and collectors under the 
name: C. nudicaulis, whose achenes are appressed, with a short but distinct 
attenuation at the apex, and all the pappus hairs are similar in thickness. 
Among other authors and collectors C. nudicaulis is understood as a 
biennial or perennial plant with prismatic or columnar achenes, close to 
those of Rhabdotheca Cass.; the pappus hairs are of differing thickness, 
typical also of Rhabdotheca. It is difficult to infer from Lessing's brief 
description which particular species of the two was taken by him as the 
basis of the genus Microrhynchus. It is even more problematic to determine 
which of them corresponds to the true Chondrilla nudicaulis. For this, it 
is essential to study the achenes of the authentic Linnaean specimen. One 
thing is clear: if C. nudicaulis is an annual plant with achenes and pappus, 
as referred to earlier, then the genus Microrhynchus has a right to 



237 

independent status (but this may threaten the status of the later estabhshed 
genus Brachyrhamphus DC). If, however, the type species is the same 
plant as the type species of Rhabdotheca, then the genus Microrhynchus 
becomes redundant. In both cases, the species widely known as Launaea 
(Microrhynchus) fallax does not agree with the diagnosis of Microrhynchus 
in the nature of the achenes and should be split off into a new genus, 
which I have named Paramicrorhynchus. Morphologically it is close to 
Lomatolepis Cass. 

1. P. procumbens (Roxb.) Kirp. comb. nova. — Prenanthes 
procumbens Roxb. Fl. Indica III (1832) 404.— Microrhynchus fallax Jaub. 
and Sp. 111. pi. or. Ill (1847-1850) 106; O. and B. Fedtsch. Perech. Rast. 
Turk. IV, 347 (Consp. Fl. Turk. No. 2846); Vvedensky et al. Opred. 
Rast. Golodnoi Stepi 40 (Tr. Tashkent. Gos. Univ. Vyp. 178, Bot.); 
238 Kovalevskaja in Fl. Uzb. VI, 479. — Zollikoferia leucodon Fisch. and Mey. 
Ex Kar. in Bull. Soc. Nat. Mosc. XII, 2 (1839) 161, nomen (Enum. pi. in 
Turcom. et Pers. bor. leg. No. 526). — Z. fallax Boiss. Fl. or. Ill (1875) 
2,24.— Launay a fallax Kuntze, Rev. Gen. I (1891) 350; Kitam. in Res. 
Kyoto Univ. Exp. Karak. a. Hinduk. 1955, II, 431 (Fl. of Afghanistan, 
I960).— Launaea fallax Muschl. A. Man. fl. Egypt. II (1912) 1060; Nevski 
in Som. Rast. SSSR IV, 345; Popov in Tr. Uzb. Gos. Univ. Nov. Ser. 
No. 27, Biol. Vyp. 14, 101; Kirp. in Fl. Turkm. VII, 332; Zakirov, Fl. i 
Rast. Bass. Zeravsh. II, 426. — L. fallax Jaub. and Sp. ex B. Fedtsch. Rast. 
Turk. (1915) 764. — L. procumbens Amin in Tackhohn, Students fl. of 
Egypt (1956) 84 (nom. illegit., sine basionymo). — Microrhynchus 
nudicaulis auct. fl. Ross. pier. As. Med., non Less. (incl. P divaricatus 
DC); Ldb. Fl. Ross. II, 2, 832; and B. Fedtsch. Perech. Rast. Turk. IV, 
347 (Consp. Fl. Turk. No. 2S45).— Launaea nudicaulis auct. fl. As. Med., 
non Hook, f.: B. Fedtsch. Rast. Turk., 764. — Ic: Jaub. and Sp. op. cit. t. 
276; Kovalevskaja op. cit. Plate XLVI; Fig. 5 (utroque casu sub 
Microrhyncho fallaci). — ^Exs.: Sintenis, It. transcasp.-pers. 1900-1901, No. 
2001 (sub Zollikoferia fallaci). 

Perennial (or biennial?). Stems few, 10-30(50) cm high, ascending 
or procumbent, weakly pubescent or almost glabrous, weakly sulcate, 
slender (usually 1-1.5 mm thick), sometimes weakly geniculate, often 
leafless (virgate), almost dichotomously branched. Lower leaves in basal 
rosette, pinnatilobate, or pinnatisect, or runcinate, with white, cartilaginous, 
usually uncinate spinules along margin; cauline leaves few, small, almost 
entire to erose-toothed and pinnatisect. Capitula cylindrical (sometimes 
slightly broadened above or with slight constriction in middle), with 15- 
20(30) florets, solitary or in groups of few at apices of branches or on 
short peduncles in axils on main stem. Involucre (three-) four- or five- 



238 

rowed, imbricate; bracts with wide, light-colored, scarious border, with 
midrib, thickening in lower part; outer bracts very small, triangular-ovate; 
middle bracts gradually expanded; inner bracts oblong-linear. Corolla 
yellow. Achenes 2.3-2.8(3) mm long, heteromorphic; outer achenes dark 
brown or gray, with rather numerous, usually light-colored, prominent, 
longitudinal tuberculate-rugose ribs, truncate at base, deltoidly narrowed 
toward apex terminating in slender, light-colored, scarcely visible, fragile 
beak; inner achenes prismatic-columnar, usually stramineous, more or less 
not tuberculate-rugose; intermediates between these two forms also found; 
pappus about three times as long as achene, of numerous, very thin, silky 
white, slightly barbed (under lens!) hairs, basally connate into ring. 
Flowering (May)June to October. (Plate XV, Fig. 2.) 

Floodplains of rivers and gravelly terraces, mostly on saline soils; in 
239 oases; sometimes in cotton fields, mulberry plantations and in orchards. 
— Soviet Central Asia: Kyzyl-Kum, Kara-Kum, mountainous Turkmenia, 
Amu-Darya, Syr-Darya, Pamiro-Alai. General distribution: western 
Mediterranean, eastern Mediterranean, Iran Region (Iran and Afghanistan), 
Indo-Himalayas, Dzhungaria-Kashgaria (Kashgaria). 

Note. Given that Microrhynchus nudicaulis is excluded from the flora 
of the USSR, the present treatment does not include the many synonyms 
of this species cited in the works of Russian taxonomists. A complete list 
can be found in Ledebour (Ldb. op. cit.). 

GENUS 1649. RhabdothecaCass.'^ 

Cass, in Diet. sc. nat. XL VIII (1827) 424. — Microrhynchus Less. §1. 
Rhabdotheca DC. Prodr. VII, 1 (1838) 180; Benth. in Benth. and Hook, f., 
Gen. pi. II (1873) 518. — Sonchus L. subgen. Rhabdotheca (Cass.) Jaub. 
and Sp. Ill pi. or. Ill (1847-1850) 110. — Launaea s. 1. auct. p. min. p. 
(non Cass, in Diet. sc. nat. XXV (1822) 321): O. Hoffm. in Pflanzenfam. 
IV, 5 (1894) 310.—Zollikoferia DC. op. cit. p. 183, p. p. (non Zollikoferia 
Nees, 1825). — Zollikoferia DC. sect. I. Euzollikoferia Boiss: Fl. or. III- 
(1875) 822, p. max. p. 

Capitula homogamous, usually with 15-35 flowers, borne singly at 
apices of branches, not drooping off at fruit maturity, cylindrical, turbinate, 
narrowly campanulate, or ovoid. Involucral bracts four- or five-rowed, 
imbricate. Receptacle flat or weakly convex, glabrous, with papilliform 
projections at point of attachment of ovary (always?). Anthers with 



' Treatment by M.E Kirpicznikov. 

- From the Greek words rhabdos — rod, twig, and thece — storehouse; the name alludes 
to the rod-shaped fruits, containing the achenes in a sheath. 



239 

sagittate appressed appendages. Style in upper part and its branches 
(outside) densely covered with short and fine, upward-directed hairs. 
Achenes columnar (rod-shaped or narrowly cylindrical according to some 
authors), with four or five strongly developed longitudinal ribs, 
heteromorphic: outer (peripheral) achenes sometimes more or less curved, 
usually five-ribbed, densely pubescent (under lens!), usually grayish, dark 
gray, ox dark brown; more or less glabrous toward center of capitulum 
and light colored, number of ribs often four, thus innermost achenes more 
or less glabrous, pale brown or stramineous; achene slightly broadened at 
base and with depressed bottom. Pappus persistent, of fewer relatively 
thick, long and erect hairs and more numerous, very soft, thin, silky, 
more or less flexuous hairs. 

Type of genus: Rhabdotheca chondrilloides (Desf.) Webb. (= Sonchus 
chondrilloides Desf., Rhabdotheca sonchoides Cass., Zollikoferia 
chondrilloides DC). 
240 The genus includes 7-10 species, of which only one is known in the 

USSR, almost exclusively on the Ust-Urt Plateau and in the Turan lowland; 
most of the species are found in northern Africa (Morocco, Algeria, Libya, 
Egypt, Lebanon, Syria, Saudi Arabia), as well as in the Pyrenees (mainly 
in the southeastern part). 

1. R. korovinii (M. Pop.) Kirp. in Fl. Turkm. VII (1960) 327; 
Kovalevskaja in Fl. Uzb. VI, 479. — Zollikoferia korovinii M. Pop. in Bot. 
Mat. Gerb. Glavn. Bot. Sada RSFSR V (1924) 37 .—Reichardia korovinii 
M. Pop. ibid, in synon. — Launaea korovinii M. Pop ex pavl. Fl. Tsentr. 
Kazakhst. Ill (1938) 360; Zakirov, Fl. i Rast. Bass. Zeravsh. II, 426— 
Ic: Kirp. op. cit. 329; Plate LI; Kovalevskaja, op. cit. Plate XLV, Fig. 1. 

Annual? Roots slender, simple. Stems 5(12)-25(40) cm high, very 
finely tomentose (sometimes mixed with isolated straight hairs) or almost 
glabrous, usually divaricately branched or divaricately forked from middle, 
only rarely weakly branched (in small plants); branches strong; terminal 
branches with very fine scale leaves, bearing solitary capitula at apices. 
Leaves somewhat thick, usually lanceolate, gradually strongly reduced 
upward, erose-sinuate to deeply pinnatisect, with whitish, dense, 
cartilaginous spinules or spinescent teeth along margin; basal leaves basally 
wide, petiolate, often crowded in rosettes; cauline leaves sessile, 
semiamplexicaul, auriculate. Capitula more or less cylindrical or narrowly 
ovoid to conical, with 15-20(35) florets. Involucral bracts usually decurved 
after fruiting, finely pubescent outside, sometimes violet; outermost bracts 
small, obovate, auriculate, usually abruptly narrowed into subobtuse more 
or less recurved cusp; middle bracts oblong-linear, with similar apex; 
innermost bracts oblong-linear, several times as long as outermost, 
subobtuse or obtuse, with wide scarious border. Corolla yellow(?). Achenes 



240 

about 10 mm long and up to 1 mm wide, columnar (narrowly cylindrical), 
with more or less deep, four or five, longitudinal furrows, slightly narrowed 
above, weakly expanded below, witii distinctly depressed hilum continued 
inward, and few radial septae convergent at center; outer achenes covered 
with very short, upward-directed, membranous appendages ,or papillose 
241 hairs (under a hand lens!), forming transverse undulate rows on achenes; 
inner achenes subglabrous or glabrous; pappus persistent, slightly shorter 
than achenes, of white heteromorphic hairs; in inner achenes five to nine, 
thicker, long, dense, slightly (especially at apex) barbed (under a hand 
lens!); in outer achenes more numerous and shorter, silky, very thin, soft, 
more or less flexuous, almost completely smooth. Flowering May to July. 
(Plate XV, Fig. 3.) 

On low hummocky or gravelly sands in desert, less often on small 
clay hills. Soviet Central Asia: Aralo-Caspian, Kyzyl-Kum, Kara-Kum, 
mountainous Turkmenia, Amu-Darya, Syr-Darya (former Kokand District). 
Endemic. Described from Turkmenia and Uzbekistan. 

As a lectotype, I propose to select the foUowig specimen, preserved 
in the V.L. Komarov Botanical Institute, Academy of Sciences of the 
USSR, Leningrad: "Fergana Region, Kokand District, sands near Kyzyl- 
Tol, southeast of village Dauduk, 23. VI. 1913, No. 882, Z.A. Minkwitz." 
The specimen has an additional label in the hand of M.G. Popov: 
"Launaea korovinii M. Pop. (sp. n.)." 

GENUS 1650. Atalanthus D. Don'^ 

D. Don in Edinb. New Phil. Joum. VI (1829) 311; Pomel, Nouv. 
mat. fl. Atl. fasc. I (1874) 7, p. min. p. — Sonchus L. sect. III. Atalanthus 
DC. Prodr. VII, 1 (1838) 189, p. p.— Sonchus L. subgen. II. 
Acanthosonchus Sch. Bip. in Parker- Webb, and Berth. Hist. nat. il. Canar. 
Ill, p. II, sect. 2 (1850) 427. 

Capitula homogamous, with 5-12 florets, usually cylindrical, solitary 
or, less often, in groups of two to a few, at apices or in axils of short 
spinose branches. Involucre three- to five-rowed, imbricate; bracts usually 
squarrose, innermost linear, oblong-linear, or elliptical, scarious along 
margin. Achenes heteromorphic: prismatic and almost not compressed, 
oblong, fusiform, or elliptical and more or less compressed with few 
(usually four) prominent ribs, or ribs many, not so prominent. Pappus of 
numerous, thin, white, silky, weakly barbed (under lens!) somewhat 
unequal hairs. 



' Treatment by M.E Kirpicznikov. 

^ From the Greek words, atalos — tender, and anthos — flower. 



241 

Lectotype of genus: Atalanthus spinosus (Forsk.) Pomel. 

Divaricately branched semishrubs or shrubs; branches usually spinose. 
The genus comprises few species, mainly found in arid areas of the 
Mediterranean Region, on the Canary Islands, in Iran, Afghanistan, 
Mongolia, northern China; Turkmenian SSR in the USSR. 

1 . A. acanthodes (Boiss.) Kirp. comb. nova. — Zollikoferia acanthodes 
Boiss. Fl. or. Ill (1875) 827; O. and B. Fedtsch. Perech. Rast. Turk. IV, 
347 (Consp. Fl. Turk. No. 2847). — Launaya acanthodes Kuntze, Rev. 
Gen. I (1891) 350. — Launaea acanthodes T. Durand and B.D. Jackson in 
242 Ind. Kew. Suppl. I, fasc. 3 (1903) 240. L. acanthodes Boiss. ex B. Fedtsch. 
Rast. Turk. (1915) 763. — L. acanthodes B.D. Jackson ex Kirp. in Fl. 
Turkm. VII (1960) 324.— Ic: Kirp. op. cit. 325, t. L (sub. Launaea 
acanthode). 

Perennial or cultivated. Stem 30-50 cm high, shallowly sulcate, 
glaucescent, very finely tomentose or glabrous, up to 5 mm thick at base, 
gradually becoming thinner above, often forked, with terminal, stiff, 
spinescent branches. Leaves withering early (as a result, stems of fertile 
plants usually leafless, virgate), stiff, glaucous, elliptical, oblong or oblong- 
spatulate, narrowed into wide petiole, erose-sinuate, with sparse, thick, 
cartilaginous spines. Capitula cylindrical or narrowly obconical, with 5- 
6(10) florets, solitary, less often two to several, on branches (often in their 
axils) on short peduncles with finely pubescent, scaly leaves, somewhat 
gradually merging with outer involucral bracts. Involucre (three-) 
four- or five-rowed, loosely pubescent on dorsal side, weakly divergent; 
outer bracts shorter, triangular or widely ovate, inner ones longer, elliptical 
or oblong-lanceolate, with wide transparent border. Corolla yellow. 
Achenes about 4 mm long, heteromorphic, some prismatic or more or 
less fusiform, glabrous or weakly pubescent, with few (often four) ribs at 
corners, obtusely truncate at base; others more or less asymmetric, 
puberulent (strongly enlarged!), with many longitudinal ribs, usually 
narrowed below; besides these, intermediates of achenes also found, with 
truncate apex forming almost squarish wide areole-bearing pappus of 
numerous thin, white, silky, weakly barbed (under a hand lens!), about 3 
mm long hairs. Flowering April to June. (Plate XV, Fig. 4.) 

Stony, gravelly and rubbly soils in deserts and foothills. — Soviet 
Central Asia: mountainous Turkmenia. General distribution: Iran. 
Described from Iran. Type in Geneva. 

Note. A quite polymorphic and inadequately studied species. 
Collections of specimens with indisputably mature achenes, as well as 
good specimens from which to derive more accurate information about 
the vegetative plant parts (especially about the root system and leaves), 
are desired. 



242 




243 Plate XV. 

1. — Sonchus transcaspicus Nevski, habit and achene; 2. — Paramicrorhynchus procumbens 

(Roxb.) Kirp.: a — outer achene, b — inner achene; 3 — Rhabdotheca korovinii (M. Pop.) Kirp. 

a — outer achene, b — inner achene; 4 — Atalanthus acathodes (Boiss.) Kirp., achenes. 



243 
244 GENUS 1 65 1 . Sonchus L. ' -^ 

L. Sp. pi. (1753) 793; DC. Prodr. VII, 1 (1838) 184, p. p.; Benth. in 
Benth. and Hook. f. Gen. pi. II (1873) 528, p. p.; O. Hoffm. in 
Pflanzenfam. IV, 5 (1894) 371, p. p.; Boulos in Bot. Notis. CXIII, 4 
(1960) 400. 

Capitula homogamous, many-flowered (each with 75-250 florets) or, 
less often, few-flowered, in corymbose, corymbose-paniculate, or 
umbellate-corymbose inflorescence. Involucre obovoid, campanulate, bowl- 
shaped or cyUndrical, often swollen below after flowering and then pitcher- 
shaped, two- or three- (four)- rowed, imbricate; involucral bracts 
herbaceous, lanceolate, oblong or narrow-triangular, usually scarious along 
margin; inner bracts two times or more as long as outermost. Receptacle 
flat. Anthers with short sagittate basal appendage. Corolla yellow. Achenes 
ovoid, oblong, oblong-ovoid or obovoid, flattened to flat, less often 
prismatic, with several (up to 20) longitudinal ribs, often transversely 
rugose. Pappus of numerous white, fragile hairs falling singly or more or 
less distinctly clustered in groups of few, or pappus hairs basally connate 
into ring easily detaching from achenes. 

Lectotype of genus: Sonchus oleraceus L. 

The genus is represented by annual, biennial, or perennial, mainly 
herbaceous plants, growing in the wild in Eurasia and Africa; several 
weedy species have been dispersed by man over almost the whole world. 
The genus, in a broader sense, comprises about 50 species. 

1. Perennials 2. 

+ Annuals (sometimes biennial?) 5. 

2. Plants with vertical root and more or less horizontal long, cord- 
like, rhizomatous. fragile lateral roots, bearing numerous 
adventitious buds. Achenes dark brown. Widespread weeds 

4. S. arvensis L, 

+ Plants with rhizomes. Achenes light-colored 3. 

3. Leaves usually 0.6-1.5 cm wide and 8-15 cm long, linear, 
lanceolate or oblong-lanceolate, undivided or, sometimes, sinuate, 
glaucous or glaucous-green. Species found in Southern 
Transcaucasia and Soviet Central Asia 

1. S. transcaspicus Nevski 



' Treatment by M.E. Kirpicznikov. 

^ The name is found in the works of ancient authors: Theophrastus used it to designate 
some thistle-like plant, while Pliny, in all probability, used it for the present Sonchus 
oleraceus L. 



244 

245 + Leaves wider, at least some of them more or less strongly divided 

4. 

4. Robust, herbaceous, usually 80-200 cm-high plants, with numerous 

long, glandular hairs in upper part and on peduncles 

2. S. palustris L. 

+ Less robust plants, usually 50-70 cm high, without glandular-hairy 

peduncles and involucral bracts 3. S. sosnowskyi Schchian 

5(1) Achenes almost always with only three filiform longitudinal ribs 

on each side, smooth and not transversely rugose between them, 

or weakly rugose, or edges of achenes usually winged and barbed 

(under lens!) 6. S. asper (L.) Hill 

+ Achenes with three or more strongly projecting longitudinal ribs 

on each side, distinctly transversely rugose, not winged 

5. S. oleraceus L. 

Section 1. Sonchus. — Sonchus sect. II. Sonchi genuini W.D. Koch, 
Synops Fl. Germ, et Helvet. (1837) 433. — Sonchus sect. Eusonchus DC. 
Prodr. VII, 1 (1838) 184. — Sonchus subgen. Sonchus: cfr. Boulos in Bot. 
Notis. CXIII, 4 (1960) 412. — Capitula many-flowered; achenes without 
beak, usually truncate at apex. Mosdy polymorphic plants with alternate 
leaves sometimes crowded in basal part of stem, usually distinctly divided, 
mosdy echinate-toothed along margin. 

Type of section: type of genus. 

Series 1. Maritimi Kirp. — Achenes stramineous or light-brown, more 
or less distinctly three- or four-angled, with few prominent ribs on each 
side, middle rib usually thicker than others. Plants often growing on saline 
soils along seacoasts as well as in arid, usually sandy habitats near water 
sources (rivers, canals, etc.). The series is named after S. maritimus L., 
which is not found in the USSR. 

1. S. transcaspicus Nevski in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 
1, 4 (1937) 293; in Somye Rast. SSSR IV (1935) 336, descr. ross.; 
Schchian in Zam. po Sist. i Geogr. Rast. Tbil. 15, 66, Kirp. in Fl. Turkm. 
VII, 332; Askerova in Fl. Azerb. VIII, 548; Kovalevskaja in Fl. Uzb. VI, 
482. — 5. maritimus auct. non L.: Boiss. Fl. or. Ill, 797; Rech. f. in Dan. 
Biol. Skr. VIII, 2, 202 (Symb. Afghanicae II); Kitam. in Res. Kyoto Univ. 
Exp. Karak. a. Hinduk. 1955, II, 448 (Fl. of Afghanistan, I960).— 5. 
baburi M. Pop. in Tr. Uzb. Gos. Univ. Nov. Ser. No. 27, Biol. Vyp. 14 
(1941) 106 (descr. ross.). — S. arvensis auct. non L.: Grigorev, Opred. 
Rast. Okr. (Dushanbe), 288.— Ic: Kovalevskaja, op. cit. Plate XLVII, 

246 Fig. 4. — Exs.: Kotschy, PI. Pers. austr. 1845, No. 109; Bomm. It. pers.- 



245 

turc. 1892-1893, No. 4112; Sintenis, It. transcasp.-pers. 1900-1901, Nos. 
276, 1648 (omnia sub 5. maritimo L.). 

Perennial. Plants with creeping rhizomes; stems (6)25-50(80) cm high, 
weakly sulcate, smooth or arachnoid-tomentose above (especially below 
capitula). Leaves glaucous or glaucous-green, narrowly linear, linear, 
lanceolate or oblong-lanceolate, (2)8-15(22) cm long and (0.2)0.6-1.5(3) 
cm wide, undivided or, less often, more or less sinuate, with short 
inconspicuous spinules along margin; lower cauline leaves narrowed 
toward base into narrowly winged, semiamplexicaul petiole; upper leaves 
sessile, with auriculate or sagittate base. Capitula two to five (sometimes 
more), on long peduncles, with about 100 florets, aggregated in lax 
corymbose inflorescence. Involucre campanulate or bowl-shaped, 8-12 
mm long, (7)10-15(20) mm wide, two- or three-rowed, basally pubescent; 
outer involucral bracts lanceolate or narrowly triangular, inner bracts 
oblong-lanceolate, with membranous border, acute or subobtuse, sometimes 
with tuft of hyaline hairs (under lens!). Corolla yellow or pale yellow. 
Achenes ovoid, oblong or (less often) oblong-ovoid, (2)2.5-3 mm long 
and 1-1.25 mm wide, usually straw-yellow, more or less distinctly three- 
or four-angled or flattened, with several (often five) prominent longitudinal 
ribs on each side; central achenes often thicker than others, with short, 
annular, reddish-brown ridge at apex; pappus of thin, soft, usually 
absolutely smooth, readily detached hairs, about two times as long as 
achene. Flowering June to September. (Plate XV, Fig. 1.) 

Foothills up to middle mountain zone; in wet places in river valleys 
and along streams. — Caucasus: Southern Transcaucasia; Soviet Central 
Asia: mountainous Turkmenia, Amu-Darya, Syr-Darya, Pamiro-Alai. 
General distribution: eastern Mediterranean, ?Armenia and Kurdistan, Iran 
Region (Iran and Afghanistan), Indo-Himalayas (Pakistan; published 
report). Described from Soviet Central Asia (Kugitang). Type in Leningrad. 

Note. Very close to the Mediterranean species S. maritimusX-, but 
differs from it by having glaucous, less serrate, almost entire leaves and 
smaller capitula. 

Some authors (cf. Zakirov, Fl. i. Rast. Bass. Zeravsh. II (1961) p. 
427) include S. transcaspicus in the synonyms of S. maritimus a 
angustifolius Bisch. (Beitr. z. Fl. Deutschl., Cichorieen (1851) 225). In 
fact, this variety was established for European specimens having no 
relationship to the Mediterranean material. 

Series 2. Palustres Schchian in Zam. po Sist. i Geogr. Rast. Tbilisi 
15 (1949) 63, p. p. — Achenes light-yellow or brownish, more or less 
distinctly tetrahedral, weakly rugose transversely. Plants of marshy and 
moist places. 



246 

247 2. S. palustris L. Sp. pi. (1753) 793 {"paluster" auct mult.); DC. 
Prodr. VII, 1, 187; Ldb. Fl. Ross. II, 2, 836; Boiss. Fl. or. Ill, 798 
Schmalh. Fl. II, 164; Fedtsch. and Fler. Fl. Evrop. Ross. 1056; O. and B 
Fedtsch. Perech. Rast. Turk. IV, 348 (Consp. Fl. Turk. No. 2851); Grossh 
Fl. Kavk. IV (1934) 254 and Opred. Rast. Kavk. 515; Ling in Contrib 
Inst. Bot. Nat. Acad. Peip. Ill, 4, 184; Krasch. in Fl. Yugo-Vost. VI, 467 
Pavlov, Fl. Tsentr. Kazakhst. Ill, 363; Kryl. Fl. Zap. Sib. XI, 3020; Klok 
in Vizn. Rosl. URSR, 602; Schchian in Fl. Gruzii VIII, 677; Szafer, Kulcz. 
B. Pawl. Rosl. Polskie (1953) 742; Kirp. in Majevski, Fl. (1954) 632 
Hayek in Hegi, 111. Fl. 2, Aufl. VI, 2, 1102; Dostal, Klic (1954) 804 
Stank, and Tal. Opred. Vyssh. Rast. (1957) 441; Mikhailovskaja in Fl 
BSSR, V, 190; Kirp in Fl.-Turkm. VII, 332; Nikitina, Mater, po. Fl. Sev 
Ski. Kirg. Ala-too, 111, No. 1132; Askerova in Fl. Azerb. VIII, 548 
Kovalevskaja in Fl. Uzb. VI, 482. S. sexpedalis Gilib. Fl. Lithuan. Ill 
(1781) 241; Ej. Exerc. phytol.I (1792) 192.— 5. paludosus Gueldenst. ex 
Ldb. Fl. Ross. II, 2 (1845-1846) 836. —S. inundatus. M. Pop. in Tr. 
Uzb. Gos. Univ., Nov. Sdr. No. 27, Bioi. Vyp. 14 (1941) 105 (descr. 
ross.); Zakirov, Fl. i Rastit. Bass. Zeravsh. II, 427. — Ic: Rchb. f.-Ic. fl. 
Germ, et Helvet, XIX (1859), t. 63 (MCCCCXIV); Syreistsch. 111. Fl. 
Mosk. Gub. Ill (1910) 335; Javorka and Csapody, Iconogr. fl. Hungar. 
(1933) t. 559, fig. 4019; Hegi, op. cit. 1102, fig. 782; Dostal, op. cit. 
805, fig. 2225; Stank, and Tal. op. cit. 442, Fig. 413.— Exs.: Rchb. Fl. 
Germ. exs. No. 1666; Fl. Gall, et Germ. exs. No. 889; Billot, Fl. Gall, et 
Germ. exs. No. 3429; Sintenis, It. transcasp.-pers. 1900-1901, No. 954. 

Perennial. Rhizome short, thick (up to 4-5 cm), producing string- 
like roots. Stems (30)80-200 cm high (according to published reports up 
to 425 cm), at base 1.5-3 cm thick, fistular, hard, sulcate, glabrous below, 
with numerous long glandular hairs above, including peduncles (only very 
rarely more or less glabrous). Leaves sessile, sagittate at base, spinose- 
toothed along margin; lower leaves large (15-35 cm long and 5-20 cm 
wide), pinnatipartite or runcinate, with few lateral triangular lobes and 
large (attenuate) apical triangular or lanceolate lobe; middle cauline leaves 
shorter than lower, lanceolate long-acuminate, undivided, spinulose-toothed 
or almost entire, uppermost leaves linear-lanceolate or linear, small, entire. 
Flowering shoots hard, usually long, slightly thickened below, bearing 
more or less large number of compact peduncles of variable length, usually 
umbellate. Capitula in corymbose or corymbose-paniculate inflorescence, 
numerous, each well-developed capitulum comprising about 75 florets. 
Involucre narrowly or broadly campanulate, 10-15 mm long; involucral 
bracts two- or three-rowed, dirty-dark green, lanceolate, densely covered 

248 on outer side (like peduncles and flowering shoots) with long glandular 
hairs. Corolla yellow. Achenes light yellow or brownish, 4-5 mm long 
and about 1 mm wide, almost tetrahedral (prismatic), slightly compressed. 



247 

weakly finely-rugose transversely, with four relatively thick lateral ribs 
and two or three not so distinct longitudinal ribs between them; achenes 
apically truncate with flat, level, quadrangular or roundish-rectangular disk 
with weakly raised (under high magnification!) reddish margins, slightly 
narrowed toward base; pappus of white or sometimes yellowish-white, 
very thin, soft, somewhat barbed (under high magnification!) rather fragile 
bristles, about two times as long as achene. Flowering May to September. 
(Plate XVI, Fig. 2.) 

Wet marshy places, riverbanks, in swamps among shrubs; in the zone 
of irrigated agriculture — along streams and impoundments. — European 
Part: (everywhere quite rare): Dvina-Pechora (southwest; only, one sheet 
seen from vicinity of Vologda), Baltic Region (published reports). Upper 
Volga, Volga-Kama (southern half), Upper Dnieper, Middle Dnieper, 
Volga-Don, Trans- Volga, ?Upper Dniester, Bessarabia (published reports), 
Black Sea Region, Crimea (one sheet seen from southern slope of Chatyr- 
Dag), Lower Don, Lower Volga; Caucasus: all regions (but rare); Western 
Siberia: all regions (but everywhere very rare); Soviet Central Asia: Aralo- 
Caspian, Balkhash, Dzhungaria-Tarbagatai, mountainous Turkmenia, Syr- 
Darya, Pamiro-Alai, Tien Shan. General distribution: Scandinavia, central 
Europe, Atlantic Europe, western Mediterranean, Balkans-Asia Minor 
(Balkan Peninsula), Dzhungaria-Kashgaria. Described from Europe. Type 
in London. 

Note. S. palustris, unlike other common and widespread Sonchus 
species, is not a weed, and probably, therefore, it is relatively less 
polymorphic. 

3. S. sosnowskyi Schchian in Zam. po Sist. i Geogr. Rast. Tbil. 15 
(1949) 72. Ic: Schchian, op. cit. t. post pag. 74, fig. 7 (solum achenium). 

Perennial. Stem 50-70 cm high, glabrous, sulcate. Leaves glabrous, 
glaucous-green, sinuate-runcinate, with fairly large, wide triangular, 
apically attenuate terminal lobe and smaller, retrorse lateral lobes, spinose- 
toothed, along margin, and with acute auricles at base. Capitula on smooth 
peduncles, medium, up to 2 cm wide, in lax, divaricate-corymbose 
inflorescence. Involucral bracts smooth; outer bracts linear-lanceolate, 
shorter than inner, lanceolate bracts, apically brownish. Achenes brownish, 
oblong-ovoid, weakly tetrahedral, with thin, almost ^cute edges and 
inconspicuous ribs in between, indistinctly rugose transversely or almost 
smooth. Flowering July. (Plate XVI, Fig. 5.) 

In wet places. — Caucasus: Southern Transcaucasia. Endemic. 
Described from the collection of A. Schelkovnik and E. Kara-Murza from 
249 the Gyunei bank of Lake Sevan. Type in Tbilisi. 

Note. The description is taken from A. Schchian, as I saw only the 
one, albeit type specimen, a poor example, without the stem base and 



248 

roots, and having hardly any capitula. This species needs greater attention; 
it is necessary to confirm its separate status with new material. 

Series 3. Arvenses Schchian in Zam. po Sist. i. Geogr. Rast. Tbilisi, 
15 (1949) 63. — Achenes light brown or dark brown, usually with five 
transversely rugose ribs on each side. Primarily a weed, widely dispersed 
by man. 

4. L. arvensis L. Sp. pi. (1753) 793; DC. Prodr. VII, 1, 187; Ldb. Fl. 
Ross. II, 2, 835; Schmalh. Fl. II, 163; Kom. in Tr. Peterb. Bot. Sada, 
XXV, 775 (Fl. Man'ch. Ill, No. 1644); Fedtsch. and Fler. Fl. Evrop. 
Ross. 1056; O. and B. Fedtsch. Perech. Rast. Turk. IV, 346; Kom. and 
Mis. Opred. Rast. Dalnevost. Kr. II 1097; Grossh. Fl. Kavk. IV (1934) 
254 and Opred. Rast. Kavk. 515; Nevski in Som. Rast. SSSR IV, 334; 
Krasch. in Fl. Yugo-Vost. VI (1936) 467 (incl. f wolgensis Krasch. ibid.); 
Perf. Fl. Sev. Kr. II-III, 375; Leskov, Fl. Malozem. Tundry, 103 (Tr. 
Sev. Bazy Akad. Nauk SSSR, Vyp. 2); Govorukhin, R. Urala 522; Pavlov, 
Fl. Tsentr. Kazakhst. Ill, 363; Sugaw. 111. fl. Saghal. IV, 1923; Kryl Fl. 
Zap. Sib. XI, 3018; Popov, Ocherk. Rast. i Fl. Karpat, 256; Klok. in 
Vizn. Rosl. URSR 602; Cronquist in Gleason, New Britt. a. Br. 111. fl. N. 
Un. St. a. adjac. Canada, 3, 534; Szafer, Kulcz., B. Pawl. Rogl. Polskie 
(1953) 742; Kirp. in Majevski, Fl. (1954) 631; Hayek in Hegi, 111. Fl. 
Aufl. VI, 2,1105; Dostal, Klic (1954) 804; Grubov, Konsp. Fl. MNR 
282, No. 1853 (Tr. Mong. Kom. Vyp. 67); Kitam. in Mem. Coll. Sc. 
Univ. Kyoto, ser. B, XXIII, 1, 147 (Compos. Jap. V); Vorobev in Tr. 
Dalnevost. Fil., Ser. Bot. Ill (V), 78 (Material from Fl. Kuril' skikh 
Ostrovov, No. 992); Stank, and Tal. Opred. Vyssh. Rast. (1957) 441; 
Mikhailovskaja in Fl. BSSR V, 188; Popov, Fl. Sr. Sib. II, 787; Askerova 
in Fl. Azerb. VIII, 547. — Hieracium arvense Scop. Fl. camiol. ed. 2, II 
(1772) 110.— Sonchus hispidus Gilib. Fl. Lithuan. Ill (1781) 241; Ej. 
Exerc. phytol. I (1792) 193. — S. arvensis L. p. integrifolius Bisch. Beitr. 
z. Fl. Deutschl., Cichorieen (1851) 228. — S. arvensis L. y. hastatus 
Kauffm. Moskovsk. Fl. (1866) 292.— Ic: Rchb. f Ic: fl. Germ, et Helvet. 
XIX (1859), t. 61 (MCCCCXII), fig. 1-3; Fedtsch. and Fler. op. cit. 1056, 
Fig. 1071 (mediocr.); Javorka and Csapody, Iconogr.. Fl. Hungar. (1933), 
t. 560, fig. 4020; Nevski, op. cit. 335, Fig. 490 (bona); Perf. op. cit. 377, 
Fig. Ill; Krasch op. cit. 469, Fig. 721 (bona); Sugaw. op. cit. 1922, t. 
885; Cronquist in Glerason, op. cit. 534; Hegi, op. cit, 1105, fig. 785 
(bona) and 786-787 (ic. phot.) Dostal, op. cit. 805, fig. 2226; 
Mikhailovskaja, op. cit. 189, Plate LXIV (mediocr.).— Exs. Billot, R. Gall, 
et Germ. exs. No. 1256; Herb. Fl. Ingr. No. 365 and 365b; Fellman, PI. 
Arct. No. 144 (f maritima); Callier, Fl. Siles. exs. No: 851; Martianow, 
250 PI. Minusinensis exs. Nos. 382, 616; Degen, PI. Hung. exs. (sine numero); 



249 

Fiori and Beg. Fl. Ital. exs., ser. II No. 1595; PL Finl. exs. Nos. 999, 
1411. 

Perennial. Plants with up to 50 cm long vertical root and more or 
less horizontal, long (up to 150 cm), string-like, tuberous, fragile, lateral 
roots, covered with numerous adventitious buds. Stems (6)40-100(150) 
cm high, erect, finely sulcate, including peduncles glandular-hairy or 
glabrous above. Leaves lanceolate or oblong, 8-20(28) cm long and 1.5- 
8(10) cm wide, somewhat stiff, almost undivided or weakly sinuate to 
pinnatilobate, pinnatisect and runcinate, more or less distinctly sharply 
spinose-toothed; lower leaves narrowed toward base into winged petiole, 
middle and upper leaves sessile, amplexicaul, basally auriculate or hastate. 
Capitula of 100-200 florets, usually few (5-10, very rarely more), in 
corymbose-paniculate or umbellate-corymbose inflorescence; peduncles 
glandular-hairy or glabrous, usually much longer than capitula. Involucre 
cylindrical or campanulate, 12-20 mm long, 7-15(25) mm wide; bracts 
lanceolate or oblong, dark green or blackish, two- or three (four)-rowed, 
often dorsally glandular-hairy. Corolla bright or golden yellow. Achenes 
oblong or ovoid, more or less distinctly compressed, dark brown, 2.5- 
3.5(5) mm long and 0.7-1 mm wide, usually with five transversely rugose 
ribs on each side, apically with low (inconspicuous) circular ridge, 
somewhat narrowed toward base; pappus of numerous, thin, soft, white 
or slightly grayish, more or less basally distinctly connate hairs, 9-13 
mm long. Flowering June to October. (Plate XVI, Fig. 6.) 

In crops, especially spring crops, lowlands and weedy places, kitchen 
gardens, roadsides, embankments of canals and peat quarries; meadows, 
steppefied areas with more or less strongly saline soil, riverbanks and 
seacoasts, shrub thickets (but usually in places where natural vegetation 
is destroyed for some reason). — Arctic: Arctic Europe; European Part: 
all regions; Caucasus: Ciscaucasia, Dagestan, Western Transcaucasia 
(reports about other regions doubtful); Western Siberia: all regions; Eastern 
Siberia: all regions; Far East: Zeya-Bureya, Uda area, Ussuri, Sakhalin; 
Soviet Central Asia: Aralo-Caspian, Balkhash, Dzhungaria-Tarbagatai, 
Kyzyl-Kum, ?Kara-Kum, Syr-Darya (rare), Tien Shan. General 
distribution: Almost throughout the world (excluding South Africa and 
South America). Described from Europe. Type in London. 

Note. An extremely polymorphic species. There exist many attempts 
to reflect its plasticity through infra-specific taxa. Among such 
classifications, based mainly on the European material, I may mention 
only two, published, indeed, in the 20th century. One of them is Rouy's 
(Flore de France, IX (1905) 204). He illegitimately proposed the new 
251 species S. vulgaris Rouy and included, as two subspecies, S. arvensis L. 
and 5. maritimus L., and for S. arvensis he described four varieties and 
one sub variety (designated by Greek letters). A somewhat simpler, but 



250 

still complex, infraspecific classification of S. arvensis was published by 
Hayek in Hegi (Hegi, op. cit. p. 1 106). 

Naturally, S. arvensis cannot be expected to be homogenous over 
the large area which it occupies in the USSR. But the specimens examined 
do not permit us to distinguish morphologically and geographically well- 
defmed races. On the basis of the present study of the indigenous material 
of S. arvensis, it is sufficient to differentiate the following forms within 
the USSR. 

1. S. arvensis f. arvensis. — ^5. arvensis. a. pedunculis calycibusque 

hispidis Weinm. Elenchus plant. (1824) 361. — S. arvensis a. genuinus 
Lindem. in Bull. Soc. Nat. Mosc. XLV, I, 2 (1872) 304.— 5. arvensis. a. 
involucris pedunculisque glanduloso-pilosis Gruner in Tr. Obshch. Ispyt. 
Prir. Khar'k. Univ. VII (1873) 37.-5. arvensis f. typicus G. Beck, Fl. 
Nied. Oesterr. II, 2 (1893) 1322. — S. arvensis var. typicus Korsh. in Mem. 
Acad. Sc. Petersb. VIII ser. VII, 1 (1898) 263 (Tentam. fl. Ross. or. No. 
774).— 5. arvensis a. vulgaris Kryl. Fl. Alt. Ill (1904) 747.— Involucral 
bracts, and often also peduncles, glandular-hairy. 

This form is found only in the European, part of the USSR, in the 
south up to Crimea and Ciscaucasia. However, on occasion, it is also 
found in the Caucasus, western Siberia and elsewhere and outside the 
USSR in all European countries. North America, and so on. It is usually 
replaced by the next form in the Far North, especially along the seacoast. 

2. S. arvensis f. maritimus (Wahl.) Rupr. in Mat. k. Blizh. Pozn. 
Prozyab. Ross. Imp. II (1845) 45 (Fl. Samojed. Cisuralens. No. 180).— 
S. arvensis (3. maritimus Wahl. Fl. Suec. II (1826) 483. — S. arvensis (3. 
maritimus Weinm. Enum. stirp. (1837) 75. — 5. maritimus auct. non L.: 
Ldb. Fl. Ross. II, 2, 835. 

Leaves narrow, oblong-lanceolate, almost undivided or weakly sinuate, 
finely toothed. Capitula solitary or few, usually large. Involucral bracts 
dirty dark green to blackish, as also the peduncles, without glandular 
hairs. 

Mainly found on transported marine sands. A common plant in the 
north of the European part of the USSR (White Sea Coasts of the Kola 
Peninsula, Kanin Peninsula, coastal belt of Timan tundra, etc.). Specimens 
from Sakhalin belong to this form. 

3. S. arvensis f. glabrescens (Guenth., Grab, and Wimm.) Kirp. stat. 
nov. — S. arvensis p. glabrescens Guenth., Grab, and Wimm. Enum. stirp. 
phanerog. Siles. (1824) 127. — S. uliginosus M.B. Fl. taur.-cauc. II (1808) 
238 and III (1819) 525; Ldb. Fl. Ross. II, 2, 834; Pavlov, Fl. Tsentr. 

252 Kazakhst. Ill, 364; Schchian in Fl. Gruzii VIII, 676; Cronquist in Gleason, 
ibid. — 5. glaber Schult. Observ. bot. (1809) 162. — ? S. intermedius 



251 

Bruckn. in Scrift. naturf. Freunde Berlin (1813) 153 (non vidi!).— 5. 
arvensis (3. pedunculis calycibusque glabris Weinm. Elenchus plant. (1824) 
361. S. arvensis (3. laevipes W.D. Koch, Synops. fl. Germ, et Helvet., ed. 
2 (1844) 498. — S. arvensis var. uliginosa Trautv. in Bull. Soc. Nat. Mosc. 
XXXIX, I, 2 (1866) 388.-5. arvensis p. laevipes Boiss. Fl. or. Ill (1875) 
798. — S. arvensis var. uliginosus Korsh. ibid. — S. arvensis (3. uliginosus, 
Kryl. ibid. — S. arvensis var. (3. glaber Schult. auct. mult. fl. Ross. — S. 
ketzkhovelii Schchian in Zam. po Sist. i Geogr. Rast. Tbil. 15 (1949) 71; 
eadem in Fl. Gruzii VIII, 676 (cum t. 438). 

Involucral bracts and peduncles without glandular hairs. 

The most widespread form of S. arvensis occurring especially 
frequently from the center of the European part of the USSR and extending 
to the south (Crimea, Caucasus) and the east (Siberia, Soviet Central Asia, 
Far East). It is difficult to assign it species rank, as at least in Siberia and 
Soviet Central Asia, f. arvensis is often found growing together with f. 
glabrescens. A. Schchian {Zam. po Sist. i Geogr. Rast. Tbilisi 15, 1949) 
holds a different opinion. He suggests that f. glabrescens should be 
considered a separate species: S. uliginosus M.B. According to him, this 
species is confined to the main steppe plains of Ciscaucasia, has a "definite 
geography," and in the Caucasus, it is vicarious in relation to S. arvensis 
But in fact, the geographic integrity of this taxon is not very well 
expressed. In our opinion, we have before us a typical ecological form. 
Boulos (L. Boulos in Bot. Notis. CXIV (1961) 57-64), who worked on 
the cytotaxonomy of the genus Sonchus, also considers S. uliginosus a 
synonym of S. arvensis. 

Based on his assumption regarding S. uliginosus as a Ciscaucasian 
steppe species, A. Schchian (op. cit., p. 71) described a new species, S. 
ketzkhovelii Schchian, typical of the montane xerophyte formations of 
southern Transcaucasia. According to the description, it differs in the 
shape (clavate), color (brownish), and partly in the sculpture of the achenes 
(with indistinct transversely rugose ribs), as well as in geography 
(mountainous Armenia and Turkish Kurdistan). The type specimen, which 
we had the opportunity to study, does not entirely resemble the type 
specimen of S. uliginosus, since long glandular hairs (under a lens!) are 
visible on some involucral bracts. Specimens practically indistinguishable 
from S. ketzkhovelii with narrow subentire leaves, only 1-few capitula, 
and glabrous peduncles are at times found in the European part of the 
USSR, Siberia, and Central Asia. All this does not permit us to consider 
S. ketzkhovelii an independent species. 

Among f. glabrescens, there are specimens collected at different times 

by Korshinsky and Kosinsky from the Volga delta. They are characterized 

by thin, large (15-20 cm long and 8-10 cm wide), shallowly sinuate- 

253 dentate leaves. I.M. Krascheninnikov (op. cit., p. 468) designated them 



252 

as f. wolgensis Krasch. On the other hand, a fairly large number of 
specimens from northern Kazakhstan and Central Asian Republics differ 
by having relatively narrow and stiff glaucous leaves. 

4. S. arvensis f. brachyotus (DC.) Kirp. stat. nov. — S. brachyotus 
DC. Prodr. VII, 1 (1838) 186; Ldb. Fl. Ross. II, 2, 835; Kom. and Alis. 
Opred. Rast. Dalnevost. Kr. II, 1098; Karav. Konsb. Fl. Yakutii 178; 
Boulos in Bot. Notis. CXIV (1961) 64. — 5. brachyotus DC. \^r. foliis 
elongatis integris Kar. and Kir. in Bull. Soc. Nat. Mosc. XIV, 3 (1841) 
458 (Enum. pi. a. 1840 in reg. Alt. et confin. collect.). — S. maritimus 
auct. non L.: Turcz. in Bull. Soc. Nat. Mosc. XXI, 3 (1848) 117 
(Fl. baic.-dahur. II, No. 715; excl. sydon. — S. nitido Vill. and S. littorali 
Rchb. qui species europaeae sunt). — S. shzucinianus Turcz. ex Herd, in 
Bull. Soc. Nat. Mosc. XLIII, I, 2 (1870) 189, pro syn. (PI. Raddeanae 
Monopet. IV, No. 233). — S. arvensis ssp. brachyotus Kitam. op. cit. 148. 

DeCandolle described this form as a separate species from Altai and 
Dauria and compared it with the Mediterranean species S. maritimus L. It 
differs from typical S. arvensis mainly by its leaves, which are more or 
less rounded at the apex, often almost entire or smaU-toothed, with solitary 
teeth to weakly sinuate. Besides, achenes of f. brachyotus are usually 
slightly larger (3.5 long and 1 nmi wide, whereas in the typical form 
they are 3 mm long and up to 0.8 mm wide), and the pappus is longer 
(up to 13 mm long, it rarely exceeds 10 nmi in the typical form). 

These differences, however, are fair from being stable, and, besides, 
other forms of S. arvensis are also found within the range of f. brachyotus. 
Form brachyotus is not characteristic of the locus classicus and is mainly 
distributed in the Far East and outside the USSR in northern China, 
Mongolia, and Japan, often on saline soils. 

In some regions, S. arvensis evidently hybridizes with 5. oleraceus. 
Such hybrid forms are found, in particular, in the Novouzensk District of 
Saratov Region and the Kazlatovskii District of the Ural Region nearby. 

Economic Importance. One of the pernicious perennial weeds, 
especially strongly infesting fields of spring crops. Besides its high seed 
productivity, this plant is capable of rapid regeneration even from small 
pieces of its fragile lateral roots ("propagation roots") -Eradication of field 
sow thistles is extremely difficult. The most common control measures 
are plowing, cutting of branches before flowering, application of 
herbicides, and so on, which are to be carried out at a general high-level 
of cultural practices. Detailed description of the biology of the field sow 
thistle and its control measures can be found in the article by V.F. 
Grodzinskaya Osot polevaya. "Biologiya Sornykh Rastenii". Posobie 
Uchitelya [Field Sow Thistle: In Biology of Weeds. Handbook for 
254 teachers]. Uchpedgiz, Moscow, 1960. At the same time, the field sow 



253 

thistle has some fodder values (for details cf. Larin et al. Kormovye 
rasteniya senokosov i pastbishch SSSR [Fodder Plants for Hay-fields and 
Pastures in the USSR] Vol. III. Sel'khozgis, Moscow-Leningrad, 1956, 
p. 671). 

Series 4. Oleracei Kirp. — Achenes more or less strongly appressed, 
light- to dark-brown, often serrate along margin, usually with three 
longitudinal ribs, finely transversely rugose or smooth in-between. 

Primarily an annual weed. 

5. S. oleraceus L. Sp. pi. (1753) 794 (solum, a leavis and P; excl. y 
and 5.); Ldb. Fl. Ross. II, 833; Turcz. in Bull. Soc. Nat. Mosc. XXI, 3 
(1848) 115 (Fl. baic.-dahur. II, No. 713); Boiss. Fl:. or. Ill, 795; Schmalh. 
Fl. II, 163; Kom. in Tr. Peterb. Bot. Sada, XXV, 776 (Fl. Man chzh. Ill, 
No. 1645); Fedtsch and Fler. Fl. Evrop. Ross. 1056; O. and B. Fedtsch 
Perech. Rast. Turk. IV, 348 (Consp. Fl. Turk. No. 2848); Kom. and Alis. 
Opred. Rast. Dalnevost. Kr. II, 1097; Grossh. Fl. Kavk. (1934) 254 and 
Opred. Rast. Kavk. 515'; Nevski in Som. Rast. SSSR IV, 337; Krasch. in 
Fl. Yugo-Vost. VI, 467; Perf. Fl. Sev. Kraya II-III, 376; Govorukhin, 
H. Urala, 522; Pavlov, H. Tsentr. Kazakhst. Ill, 362; Sugaw. 111. fl. Saghal. 
IV, 1925; Kolak. Fl. Abkhazii IV, 304; Kryl. Fl. Zap. Sib. XI, 3017; 
Klok. in Vizn. Rosl. URSR, 602; Cronquist in Gleason, New Britt. a Br. 
111. fl. N. Un. St. a. adjac. Canada, 3, 534; Schchian in Fl. Gruzii VIII, 
675; Szafer, Kulcz., B. Pawl. Rosl. Polskie (1953) 742; Kirp. in Majevski, 
Fl. (1954) 631; Hayek in Hegi, 111. Fl. 2 Aufl. VI, 2, 1107; Dostal, Klic 
(1954) 804; Kitam. in Mem. Coll. Sc. Univ. Kyoto, ser. B, XXIII, 1. 146 
(Compos. Jap. V); Vorobev in Tr. Dalnevost. Fil. Ser. Bot. Ill (V), 78 
(Materialy k. Fl. Kuril, skikh 0-vov, No., 993); Stank, and Tal. Opred. 
Vyssh. Rast. (1957) 441; Karav. Konsp. Fl. Yakutii 178; Mikhailovskaja 
in Fl. BSSR V, 187; Popov, Fl. Sr. Sib. II, 788; Kirp. in Fl. Turkm. VII, 
333, Nikitina, Mater, po R. Sev. Ski. Kirg. Ala-too 111, No. 1134; Kitam. 
in Res. Kyoto Univ. Exp. Karak. a. Hinduk. 1955, II, 448 (Fl. of 
Afghanistan 1960); Askerova in Fl. Azerb. VIII, 549; Vvedensky et al. 
Opred. Rast. Golodnoi Stepi (1961) 40 (Tr. Tashk. Gos. Univ Vysh. 178, 
Bot.); Kovalevskaja in H. Uzb. VI, 480.— 5. ciliatus Lam. R. Fr. II (1778) 
87.-5. glaber var. 1 and 2. Gilib. Fl. Lithuan. Ill (1781) 242; Ej. Exerc. 
phytol. I (1792) 193. —S. levis Vill. Hist. pi. Dauph. Ill (1789) 158.-5. 
asper Hall, ex Gaertn., Mey. and Scherb. Fl. Wett. Ill, 1 (1801) 125.— 5. 
oleraceus p. M.B. Fl. taur.-cauc. n (1808) 239.-5. roseus Bess, in Spreng. 
Syst. veg. Ill (1826) 651. — 5. oleraceus var. rubrif alius Lindem. in 
Bull. Soc. Nat. Mosc. XL, I, 2 (1867) 541.— 5. oleraceus. a. genuinus 
Gruner in Tr. Obshch. Isp. Khar'k. Univ. VII (1873) 31.—Sonchus 
rotundilobus M. Pop. in Tr. Uzb. Gos. Univ. Nov. Ser. No. 27, Biol. 



254 

Vyp. 14 (1941) 106, nomen. — S. rotundifolius M. Pop. ex Vass. Som. 
Rast. Tadzh. 1 (1953) 440 (erroneo loco nominis praecedentis). — Ic: O.F. 
Muller, Fl. Danica IV (1777) t. 682 (color.); Rchb. f. Ic. fl. Germ, et 
Helvet. XIX (1858), t. 59 (MCCCCX) fig. 1; Syreistsch. 111. Fl. Mosk. 
255 Gub. Ill (1910) 333; Javorka and Csapody, Iconogr. F . Hungar. (1933), 
t. 560, fig. 4021; Sugaw. op. cit. 1924, t. 886 A; Cronquist in Gleason, 
op. cit. 534; Hegi, op. cit. t. 277, fig. 3 and 3a; Dostal, op. cit. 807, fig. 
2227.— Exs.: Billot, Fl. Gall, et Germ. exs. No. 1911; Herb. Fl. Ingr. No. 
363; Todaro, Fl. Sic. exs. No. 785; Sintenis and Rigo, It. cypr. 1880, No. 
288; Bomm. It. pers.-turc. 1892-1893, No. 4110; Wolosz. Fl. polon. exs. 
No. 845; Sintenis, It. transcasp.-pers. 1900-1901 No: 366; Fl. terr. Israel, 
exs. No. 707. 

Annual (sometimes biennial?). Root narrowly conical or fusiform. 
Stem 30-100 cm high, finely sulcate, fistular, simple or branched, glabrous 
or glandular-hairy above. Leaves not stiff, dull, 10-18(22) cm long and 
5-7(12) cm wide, pinnatipartite or lyrate with large terminal, triangular 
lobe, finely sinuate-toothed, with somewhat spinescent teeth; lower leaves 
on winged petiole with wide amplexicaul base; upper leaves sessile, with 
sagittate auricles at base. Capitula mostly few (usually 5-15), in corymbose 
or umbellate-corymbose inflorescence; peduncles and bases of capitula 
arachnoid- or flocculose-lanate when young, sometimes glandular hairy; 
some capitula often undeveloped, normally with 100-125(150) florets. 
Involucre campanulate or bowl-shaped, 10-12 mm long and (6)10-15(25) 
mm wide; bracts dark green, usually .three-rowed, sometimes dorsally 
with few glandular hairs. Corolla yellow, less often whitish. Achenes 
oblong-obovoid, somewhat narrowed toward base, more or less strongly 
compressed, light brown, brown, or dark brown, 2.5-3, mm long, 1 mm 
wide, weakly toothed along margin (under high magnification!), with three 
or less often more prominent longitudinal ribs on each side, finely rugose 
transversely; pappus of numerous thin, soft, white, almost smooth bristles 
(slightly barbed under high magnification), two times as long as achene. 
Flowering June to October. (Plate XVI, Fig. 8.) 

A weed in fields, kitchen gardens, gardens, on irrigated lands, banks 
of mountain streams, less often on coastal sands; from lowlands to middle 
mountain zone. — European Part: all regions except northern part of 
Karelia-Lapland; Caucasus: all regions; Western Siberia: 70^, Upper 
Tobol, Irtysh, Altai; Eastern Siberia: Yenisei, Lena-Kolyma (very rarely), 
Angara-Sayans, Dauria; Far East: Zeya-Bureya, Uda area, Ussuri, Sakhalin 
regions; Soviet Central Asia: Aralo-Caspian, Balkhash, Dzhungaria- 
Tarbagatai (Dzhungarian Alatau), Kyzyl-Kum, mountainous Turkmenia, 
Syr-Darya, Pamiro-Alai, Tien Shan. General distribution: almost all over 
the world. Described from Europe. Type in London. 

Note. The species is particularly notable for varying in leaf shape. 



255 

corolla color, and, less often, the number of longitudinal ribs on the 
achenes and nature of its surface. 

Usually the following forms, without definite geographic correlation, 
are distinguished in S. oleraceus on the basis of the most striking character, 
the nature of the incision of the leaves. 

256 1. S. oleraceus f. integrifolius (Wallr.) Kirp. stat. nov. — S. oleraceus. 

a. integrifolius Wallr. Sched. crit. I (1822) 432. — S. oleraceus y. 
integrifolius Bisch. Beitr. z. Fl. Deutschl., Cichorieen (1851) 221. — Leaves 
undivided or subentire. Found rarely. 

2. S. oleraceus f. triangularis (Walk.) Kirp. stat. nov. — S. oleraceus 
p. triangularis Wallr. I.e. — S. oleraceus [3. runcinatus W.D. Koch, 
Synops. fl. Germ, et Helvet. (1837) 433. — S. oleraceus var. a. runcinatus 
sub-var. triangularis Coss. and Germ. Fl. envir. Paris II (1845) 437. — S. 
oleraceus a. runcinatus Bisch. op. cit. 220. — Leaves usually pinnatipartite, 
runcinate, with large terminal conical lobe. Found generally and 
frequently. 

3. S. oleraceus f. lacerus (Wallr.) Kirp. stat. nov. — S. oleraceus y. 
lacerus Wallr. 1. c.—S. lacerus Willd. Sp. pi. Ill, 3 (1803) 1513.— 5. 
oleraceus var. (3. lacerus Coss. and Germ. 1. c. — S. oleraceus (3. lacerus 
Bisch. op. cit. 220. — Leaves pinnatifid or pinnatipartite, with sinuate- 
toothed lobes or segments; terminal lobe or segment weakly distinguishable 
from laterals. Found frequently. 

Each of these forms may be represented by populations in which 
glandular hairs are found in the upper part and sometimes also on the 
involucral bracts, or by eglandular populations. In western European 
works, sometimes forms with differences in corolla color, growth vigor, 
and other characters of secondary importance for this species are separately 
designated. 

5. oleraceus hybridizes with S. asper. Among these hybrids is included 
S. rotundilobus M. Pop. (cf. Kovalevskaja, op. cit. p. 481). 

Economic Importance. A common weed which is eradicated by 
cultivation before fruiting and also by using herbicides. Young plants are 
relished by cattle and are particularly good for feeding pigs. 

6. S. asper (L.) Hill, Herbar. Britan. I (1769) 47; Fedtsch. and Fler. 
Fl. Evrop. Ross. 1056; Kom. and Alls. Opred. Rast. Dal'nevost. Kr. II 
1097; Grossh. Fl. Kavk. IV (1934) 254 and Opred. Rast. Kavk. 515; 
Krasch. in Fl. Yugo-Vost. VI, 466; Govorukhin, Fl. Urala, 522; Takht. 
and Fed. Fl. Erevana (1945) 340; Kryl. Fl. Zap. Sib. XI 3017; Kolak. Fl. 
Abkhazii, IV, 304; Klok. in Vizn. Rosl. URSR, 602; Schchian in Fl. Gruzii 
VIII, 675; Cronquist in Gleason, New Britt. and Br. 111. Fl. N. 



256 




257 Plate XVI. 

1 — Lactuca winkleri Kirp., habit, achene; 2 — Sonchus palustris L., achene; 3 — Lactuca 

sibirica (L.) Benth. ex Maxim., achene; A — L. tatarica (L.) C.A. Mey., achene; 5 — Sonchus 

sosnowskyi Schchian, achene; 6 — S. arvensis L., achene; 7 — S. asper (L.) Hill., achene; 8 

S. oleraceus L., achene. 



257 

258 Un. St. a. adjac. Canada 3, 534; Szafer, Kulcz., B. Pawl. Rogl. Polskie 

(1953) 742; Sobolevskaya, Konsp. Fl. Tuvy 200; Kirp. in Majevski, Fl 

(1954) 631; Hayek in Hegi, 111. Fl. 2 Aufl. VI, 2 1 109; Dostal Klec (1954) 
804; Vorobev in Tr. Dal'nevost. Fil. Ser. Bot. Ill (V), 78 (Mater, po Fl. 
Kuril'skikh o-vov. No. 993); Karav. Konsp. Fl. Yakutii 178; 
Mikhailovskaja in Fl. BSSR V, 188; Kirp. in Fl. Turkm. VII, 333; 
Askerova in Fl. Azerb. VIII, 549; Vvedensky et al. Opred. Rast. Golodnoi 
Stepi (1961) 41 (Tr. Tashk. Cos. Univ. Vysh. 178, Bot.); Kovalevskaja 
in Fl. Uzb. VI, 481. 5. oleraceus; y. asper and 6. L. Sp. pi. (1753) 794.— 
S. spinosus Lam. Fl. Fr. II (1778) 86.-5. glaber var. 3. Gilib. Fl. Lithuan. 
Ill (1781) 242; Ej. Exerc. Phytol. I (1792) 193.— 5. oleraceus (3. asper 
Falk. Beitr. II (1786) 232.-5. asper Vill. Hist. pi. Dauph. Ill (1789) 
158; Ldb. Fl. Ross. II, 2, 834; Boiss. Fl. or. Ill, 796; Schmalh. Fl. II, 
163; O. and B. Fedtsch. Perech. Rast. Turk. IV, 348 (Consp. Fl. Turk. 
No. 2849); Ling in Contrib. Inst. Bot. Nat. Acad. Peip. Ill, 4, 185; Nevski 
in Som. Rast. SSSR IV, 338; Perf. Fl. Sev. Kr. II-III, 376; Pavlov Fl. 
Tsentr. Kazakhst. Ill, 363; Sugaw. 111. fl. Saghal. IV, 1925; Popov, Fl. Sr. 
Sib. II, 788; Nikitina. Mater, po Fl. Sev. Ski. Kirg. Ala-too, 111, No. 
1133.— 5. fallax Wallr. Annus Bot. (1815) 98; Ej. Sched. crit. I (1822) 
432; DC. prodr. VII, 1, 185. — S. asper Vill. b. lacerus Meinsh. Fl. Ingr. 
(1878) 188.-5. eryngiifolius Sosn. in Grossh. Fl. Kavk. IV (1934) 254 
and Opred. Rast. Kavk. 515 (descr. ross.). — S. eryngiifolius Sosn. in 
Schchian in Zam. po Sist. i Geogr. Rast. Tbil. 15 (1949) 71 (descr. lat.).— 
Ic: O.F. MuUer, Fl. Danica, V (1782), t. 843 (color.); Hill, op. cit. t. 34,- 
fig. 2; Rchb. f. Ic. fl. Germ, et Helvet. XIX (1858) t. 59 (MCCCCX), fig. 
1 and t. 60 (MCCCCXI); Syreistsch. 111. Fl. Mosk. Gub. Ill (1910) 334; 
Javorka and Csapody, Iconogr. Fl. Hungar. (1933), t. 560, fig. 4022; 
Nevski op. cit. 338, Fig. 491 (solum achenium); Sugaw. op. cit. t. 886, 
B; Cronquist in Gleasdn, op. cit. 534; Hegi, op. cit. 1 109, fig. 789 (bona); 
Dostal, op. cit. 807, fig. 2228 (solum folium).— Exs.: Sch. Bip. 
Cichoriaceotheca Suppl. II, No. 156 (sub. S. aspero Fuchs); Rchb. Fl. 
Germ. exs. No. 1665 (sub S. aspero Hall.); Billot, Fl. Gall, et Germ. exs. 
No. 1912; Herb. Fl. Ingr. No. 364; Sintenis, It. transcap.-pers. 1900-1901, 
No. 1815a; Woronow, PL Abchaz. exs. (No. 266). 

Annual. Root slender, narrowly conical or fusiform, usually branched. 
Stem (10)30-75(100) cm high, finely sulcate, fistular, simple or branched, 
glabrous or covered with dark, stalked, glandular hairs above. Leaves 
somewhat stiff or almost coriaceous, less often thin and soft, dark green 
or sometimes light bluish, oblong or obovate, 6-15 cm long and 1.5-8 
cm wide, undivided, sinuate-pinnatifid, or, less often, pinnatisect, with 
unevenly spinescent teeth; lower leaves on winged petiole, middle and 
upper leaves sessile, with roundish auricles appressed to stem. Capitula 
less numerous (usually 5-10), in corymbose or umbellate-corymbose 



258 

259 inflorescence; peduncles glabrous or glandular hairy, often much longer 
than usually undeveloped capitula; fully developed capitula with 100- 
150 florets. Involucre campanulate or bowl-shaped, 10-12(15) mm long, 
(6)10-15(25) mm wide, bracts dark green, narrowly ovate, lanceolate or 
oblong, usually three-rowed. Corolla yellow. Achenes oblong-obovoid, 
flat, light- or dark-brown, 2.5-3(3.5) mm long and 0.75-1 mm wide, not 
rugose transversely (or weakly rugose), usually winged, weakly toothed 
(under high magnification!) along margin, mostly with three prominent 
longitudinal ribs on each side; pappus of numerous thin and soft, white, 
almost smooth, weakly toothed (under high magnification), fragile hairs, 
two times or slightly more as long as achenes. Flowering June to 
September (Plate XVI, Fig. 7). 

Fields, kitchen gardens, orchards and vineyards, roadsides, weedy 
places, thickets of steppe shrubs, river valleys (in semidesert), and banks 
of canals; up to middle mountain zone. — European Part: all regions except 
northern part of Karelia-Lapland; Caucasus: All regions; Western Siberia: 
all regions; Eastern Siberia: all regions; Far East: (very rarely), Uda 
area, Ussuri (southern), Sakhalin regions; Soviet Central Asia: ?Aralo- 
Caspian (western part), Balkhash, Dzhungaria-Tarbagatai, Kyzyl-Kum, 
Kara-Kum (Krasnovodsk District), mountainous Turkmenia, Syr-Darya, 
Pamiro-Alai, Tien Shan. General distribution: Scandinavia, central Europe, 
Atlantic Europe, Mediterranean, Balkans-Asia Minor, Armenia and 
Kurdistan, Iran, Indo-Himalayas, Dzhungaria. Kashgaria, ?Mongolia, 
Japan, China, North America. Besides, sporadically found in Central 
America, Australia and other regions. Described from Europe. Type in 
London. 

Note. A species higly variable in the consistency and cutting of leaves. 
Two forms, which lack geographic discreteness, are well distinguished. 

1. S. asper f. inermis (Bisch.) Kirp. stat. nov. S. asper a. inermis 
Bisch. Beitr. z. Fl. Deutschl., Cichorieen (1851) 222. — Cauline leaves fairly 
thin and smooth, oblong-ovate, entire, with short, not very stiff teeth. 

2. S. asper f. pungens (Bisch.) Kirp. stat. nov. — S. asper p. pungens 
Bisch. 1. c. — Cauline leaves more rough, often stiff or more or less 
coriaceous, deeply sinuate-toothed to sinuate or pinnatifid, with unequal, 
usually long, stiff, spinescent teeth. 

Within these varieties, western European authors often further 
distinguish individual forms depending on the color of the lamina, presence 
or absence of glandular hairs on the peduncles, and so on. 

S. eryngiifolius should be referred to 5. asper f. pungens. This species 
was described as an endemic of inner montane Dagestan, growing in the 
natural formations of montane xerophytes. But identical specimens are 



259 

260 also found in Abkhazia and outside the Caucasus. The morphology of 
S. eryngiifolius (type specimen examined!) clearly indicates that its affinity 
is closest, not to S. oleraceus as stated in the original description, but to 
S. asper. S. eryngiifolius is one of the S. oleraceus x S. asper hybrids 
without a definite localized geographic area. The traces of transverse 
wrinkles on the achenes resemble those of S. oleraceus, but other achene 
characters (presence mostly of three, fine longitudinal ribs) and the 
morphology of the vegetative organs are close to S. asper. Apparently, S. 
asper f. pungens is represented on the whole, (if not entirely) by specimens 
of hybrid origin. 

Not that long ago some authors considered S. asper as a whole as 
only a subspecies or variety of S. oleraceus (see, for example, A. Fiori, 
Nuova flora analitica D' Italia, Vol. II, Firenze, 1925-1929, pp. 861- 
817), but no one shares this viewpoint any longer. 

Kitamura (S. Kitamura in Res. Kyoto Univ. Exsp. Karak. a. Hinduk. 
1955, II (1960) 431 (Flora of Afghanistan)) cites S. asper Garsault, Fig. 
pi. et anim. Med. IV (1764) t. 565. We do not consider it necessary to 
accept the name of Garsault, because in composing it the rules of binomial 
nomenclature were inconsistently applied. 

Meyer, Turczaninow, Ruprecht and some other Russian botanists cite 
S. asper Fuchs. Although the binomial name S. asper was actually used 
for the first time by Fuchs way back in 1542, this name, being pre- 
Linnaean, is not valid. 

Economic Importance. It has little value as a fodder and food plant. 
According to some reports, it is a good honey-producing plant.. 

Doubtful and Deleted Species 

1-2. S. amplexicaulis, Gueldenst. and S. perfoliatus Gueldenst. — 
Both names were unclear already to C. Ledebour (he cited them with the 
remark "Quid?"). They also have been unclear to all subsequent workers, 
remaining unexplained nomina nuda until now. 

2. S. tenerrimus L. Sp. pi. (1753) 794. 

This Mediterranean species was wrongly reported for the Crimean 
flora by some foreign authors. 

GENUS 1652. Reichardia Roth'^ 

Roth, Bot. Abhandl. u. Beobacht. (1787) 35.— Picridium Desf. Fl. 
Atlant. II (1799) 220. 



' Treatment by M.E. Kirpic-nikov. 

2 Genus named in honor of Johann Jacob Reichard (1743-1782), a clinician and 
botanist, author of the then well-known Flora of Frankfurt-on-Main. 



260 

Capitula homogamous, many-flowered (containing up to 100 or more 
florets), terminating at apex of stem or its long branches. Involucre 
campanulate, cylindrical, or ovoid, four- or five-rowed, imbricate; bracts, 
261 herbaceous, scarious along margin; innermost bracts two to four times as 
long as outermost bracts, often with barbule of thin hairs at apex. 
Receptacle flat, more or less alveolate; corolla yellow; anthers with short, 
sagittate, appressed basal appendages. Achenes polymorphic: outer achenes 
usually prismatic or oblong, four- or five-angled and with four or five 
deep furrows, transversely tuberculate or scarious-rugose; inner achenes 
usually slightly conical, less distinctly transversely tuberculate or 
transversely rugose. Pappus falling entirely, of numerous white, unequal 
hairs, basally connate into ring. 

Lectotype of genus: Reichardia tingitana Roth. 

Annual or perennial, herbaceous, glabrous plants growing on Canary 
Islands, in the Mediterranean countries, Ethiopia, Asia Minor, Iraq, Iran, 
?Afghanistan, and the northwestern part of the Indian Peninsula. The genus 
includes in all seven or eight species, of which we have only one in the 
Caucasus. 

1. R. dichotoma (Vahl) Freyn in Oest. Bot. Zeitschr. XLII (1892) 
267; Post, Fl. or Syr., Palest, a. Sinai, ed. 2, II 152; Grossh. Fl. Kavk. IV 
(1934) 259 and Opred. Rast. Kavk. 517; Takht. and Fed. Fl. Erevana 
341; Kolak. Fl. Abkhazii IV, 306; Schchian in Fl. Gruzii VIII, 694; 
Karjagin in Fl. Azerb. VIII, 562. — Scorzonera dichotoma Vahl, Symb. 
bot. II (1791) 89. Sonchus dichotomus Willd. Sp. pi. Ill, 3 (1803) 1517; 
M.B. Fl. taur.-cauc. II, 240. — Picridum dichotomum Fisch. and Mey. ex 
DC. Prodr. VII, 1 (1838) 183; Ldb. Fl. Ross. II, 2, 832; Boiss. Fl. or. Ill, 
829. — Ic: Karjagin. op. cit. 516, Plate LVII.— Exs. Bomm. PI. exs. Anatol. 
or. 1889, No. 701; Sintenis, It. or. 1892, No. 4503. 

Perennial. Stem (30)60-100 cm high, weakly sulcate, glabrous, more 
or less strongly divaricately branched, with long, upward-directed branches, 
terminating in capitulura. Leaves flat, glaucous, oblong-linear, linear- 
lobate, or broadly lanceolate, sinuate-lobate or more or less undivided, 
spinose-toothed or spinose along margin, weakly crisped-flexuous; lower 
leaves on winged semiamplexicaul petiole, often clustered in rosette, upper 
leaves reduced, sessile, auriculate. Capitula with (35)50-70(100) florets. 
Involucre campanulate-cylindrical or campanulate, (6)10-15(20) mm long; 
involucral bracts four- or five-rowed, herbaceous, usually purple; scarious 
along margin, very finely appressed-hairy dorsally (under high 
magnification!); inner (longest) bracts subobtuse with barbules of very 
thin and short hairs (under a hand lens!). Corolla Ught yellow. Achenes 
polymorphic: outer achenes prismatic or campanulate. four-, less often 
five-angled, 4-5(7) mm long and 0.7-0.8(1.2) mm wide, dark brown. 



261 

with four or five longitudinal furrows, transversely rugose; becoming 

262 lighter in color towards centre of achene, tubercles disappearing gradually; 
innermost achenes light or stramineous, more or less smooth; pappus one 
and one-half to two times as long as achene, of almost entirely smooth, 
numerous thin hairs white or yellowish below, few among them thicker 
(under lens!). Flowering July to September. (Plate XIX, Fig. 8). 

In mountains, on dry clayey and stony slopes, in dry riverbeds. — 
Caucasus: all regions except Talysh. General distribution: eastern 
Mediterranean, Balkans-Asia Minor (Asia Minor), Armenia and Kurdistan, 
Iran (northern). Described from Tunisia. Type possibly in Copenhagen. 

Note. Grossheim and other authors of the Flora of Caucasus used 
the name "Reichardia dichotoma (M.B.) Roth." In fact, Albrecht Roth 
never used such a combination. 

Contrary to the generally held opinion, ^^Sonchus dichotomus M.B." 
should not be the basionym for R. dichotoma. Marschall-Bieberstein did 
not describe such a species; he clearly indicated that the specimens 
collected from the North Caucasus and the region of the present Tbilisi 
were included by him under 5. dichotomus Willd. 

GENUS 1653. Prenanthes hP 

L. Sp. pi. (1753) 797, p. p.; DC. Prodr. VII, 1 (1838) 194, p. p.; 
Benth. in Benth. and Hook. f. Gen. pi. (1873) 527; O. Hoffm. in 
Pflanzenfam. IV, 5 (1894) 375. 

Capitula homogamous, with variable number of flowers (5, 8-10, 15 
or 25-35 florets). Narrowly cylindrical, narrowly campanulate or very 
rarely ovoid, in lax paniculate, less often recimose- or corymbose- 
paniculate inflorescence. Involucre two- or three (four)-rowed bracts; 
herbaceous, usually dark (dirty green, dark violet, etc.) or dorsally 
pubescent (under a hand lens!) (sometimes covered with papilliform hairs) 
with isolated, erect, thin, scaly, often glandular hairs. Receptacle glabrous, 
flat. Achenes oblong-linear or linear, usually slightly compressed, less 
often almost terete, with inconspicuous longitudinal ribs (only with 
nonspecific striation), or ribs four or five, more or less well developed, 
sometimes more numerous and distinct, achenes covered with numerous, 
short, erect hairs (visible under a hand lens!), less often more or less 
glabrous, usually slightly narrowed toward base, without beak but with 

263 broad disk at apex with deciduous pappus, less often pappus persistent; 
pappus bristles numerous, dirty-rusty or light rusty, sometimes white. 



' Treatment by M.E. Kirpicznikov. 

^ From the Greek words prenes — inclined forward and anthos — flower; the name 
indicates a weak general inflorescence often inclined forward. 



262 

distinctly barbed almost throughout smooth (visible only under high 
magnification). Basic chromosome number x = 9, 8. 

Lectotype of genus: Prenanthes purpurea, L. 

Perennial herb, distributed mainly in Eurasia and North America. The 
genus comprises about 25 species. However, this number is very 
approximate, as even among the Lactuceae, the genus Prenanthes is known 
for its indistinct boundaries. Because of the present trend of combining 
Nabalus with Prenanthes, which we have also followed for the "Flora of 
the USSR, " the limits of the latter genus become even more indistinct. 

1. Corolla yellow; florets in capitulum 15-35 2. 

+ Corolla light blue, purple or violet-purple, if light colored, then 

florets in capitulum fewer 4. 

2. Florets about 15 per capitulum; laminas oblong elliptical, linear 
or linear-lanceolate. Species growing in Altai and Tuva ASSR .. 

7. P. angustifolia Boulos. 

-I- Florets 25-35 per capitulum; laminas of cauline leaves lyrately 
pinnatisect or rotund-triangular to triangular 3. 

3. Achenes 7-8 mm long, with numerous promdnent longitudinal ribs; 
pappus of dirty rusty or rusty, often barbed hairs. Plants growing 
in the Far East 5. P. maximowiczii Kirp. 

+ Achenes about 4 mm long, with four or five distinctly raised and 
indefinite number of less distinct longitudinal ribs; pappus of snow- 
white, almost entirely smooth hairs. Plants growing in the Caucasus 

6. P. abietina (Boiss.) Kirp. 

4(1) Capitula with few, usually five florets each 5. 

+ Capitula with larger number of florets 6. 

5. Lamina and petioles more or less densely covered with scaly hairs, 
and sometimes also spinulose; pappus hairs dirty-rusty or almost 

rusty. Species growing in the Far East 

2 P. tatarinowii Maxim. 

+ Lamina and petioles without scaly hairs or spinules; pappus hairs 
white. Species growing in the European part of the USSR and the 

Caucasus 1. P. purpurea L. 

6(4) Florets usually 8-10 per capitulum; leaves more or less 

crowded in lower part of stem, with relatively small lamina, 3- 

5 cm wide at base, and 8-10 cm long; stem 30-50(70) cm high, 

264 slender (usually 1-3 mm thick). Plants mainly distributed in 

western Transcaucasia 4. P. pontica (Boiss.) Leskov. 

+ Florets about 15 per capitulum. Plants more or less uniformly leafy; 
lower leaves usually much larger. Stem 50-100(150) cm high, 

thicker. Species growing almost throughout the Caucasus 

3. P. cacaliifolia (M.B.) Beauverd 



263 

Subgenus 1. Prenanthes. — Prenanthes subgen. Euprenanthes 
Babcock. Stebbins and Jenkins in Cytologia, Fujii Jubil. vol. (1937) 190, 
solum nomen. — Capitula with 5-15 florets; corolla light blue, purple, 
purple-violet, or lilac, less often pale with violet tip; achenes dark brown, 
with few prominent longitudinal ribs. Plants mainly of the Old World. 

Type of subgenus: type of genus. 

1. P. purpurea L. Sp. pi. (1753) 797; DC. Prodr. VII, 1, 194; Ldb. 
Fl. Ross. II, 2, 839; Boiss. Fl. or. Ill, 803; Schmalh. Fl. II, 170; Fedtsch. 
and Fler. Fl. Evrop. Ross. 1064; Popov, Ocherk Rast. i Fl. Karpat 256; 
Klok. in Vizn. Rosl. URSR 605; Stank, and Tal. Opred. Vys. Rast. (1957) 
444; Grossh. Fl. Kavk. IV (1934) 265 and Opred. Rast. Kavk. 519; Kolak. 
Fl. Abkhazii IV, 310; Schchian in Fl. Gruzii VIII, 709; Szafer, Kulcz., B. 
Pawl. Ro<?l. Polskie (1953) 744; Dostal, Klec (1954) 802; Hayek in Hegi, 
111. Fl. 2 Aufl. VI, 2, 1180. —P. tenuifolia L. op. cit. 797; Ldb. op. cit. 
839. — Chondrilla purpurea Lam. Fl. Fr. II (1778) 105. — C. tenuifolia 
Lam. Encycl. meth. II (1786) 78. — Prenanthes purpurea var. tenuifolia 
W.D. Koch, Taschenb. d. Deutsch. u. Schweiz. Fl. (1844) 316. —P. 
purpurea. (3. angustifolia W.D. Koch, Synops. fl. Germ, et Halvet. ed. 2, 
II (1844) 494; Boiss. Fl. or. Ill, 803; Grossh. Fl. Kavk. IV (1934) 265.— 
Ic: Rchb. f. Ic. fl. Germ, et Helv. XIX (1858), t. 57 (MCCCCVIII); 
Fedtsch. and Fler. op. cit. 1064, Fig. 1078; Bonnier, Fl. Compl. 111. France, 
Suisse et Belg. VI (1923), t. 352, fig. 1685 (color.); Javorka and Csapody, 
Iconogr. Fl. Hungar. (1933), t. 566, fig. 4061 (bona); Szafer, Kulcz., B. 
Pawl. op. cit. 745, fig. 1648; Dostal, op. cit. 803, fig. 2224; Hegi, op. cit. 
t. 279, fig.l (color, bona). — Exs.: Fl. Cauc. exs. No. 173; Baldacci, It, 
Alb. 1900, No. 183; Fl. exs. austro-hung. No. 3399; Hayek, Fl. Stir. exs. 
No. 1092; Fl. exs. reip. Bohem.-Slov. No. 276; PI. Polon. exs. No. 379. 
Perennial. Rhizome slender, woody. Stem 60-100(150) cm high, 
usually solitary, glabrous or subglabrous, paniculately branched above. 
Leaves thin, variable, usually lanceolate, oblong-elliptical or oblong-hnear, 
acute, glaucous beneath, green above; lower leaves mostly on winged 
petiole, almost entire with slightly toothed margin to sinuate-toothed and 
lyrate-pinnatipartite with large terminal lobe; middle and upper 
265 leaves sessile, semiamplexicaul, more or less entire, upper leaves smaller, 
with distinctly auriculate base; sometimes all or most leaves oblong-linear 
or linear-lanceolate to linear, entire or almost entire (var. angustifolia 
W.D. Koch). Capitula cylindrical, 12-15 mm long and 3 mm wide, 
drooping, terminal on slender branches of lax panicle, sometimes up to 
20-35 cm long; basal branches of panicle arising from axils of rather 
large upper leaves. Involucre two- or three-rowed; outer involucral bracts 
oblong-ovate, dark, with more or less dense, often papilliform pubescence 
(under a hand lens!); innermost bracts two to three times as long as outer, 



264 

linear or oblong-lanceolate, greenish in lower part, usually dark violet 
above, bordered along edges, subobtuse, with short barbules of thin, light 
colored, hairs (under a hand lens!); florets in capitulum usually five; corolla 
purple or violet-purple, its lobes covered with papilliform hairs. Achenes 
4.5-5 mm long, about 1 mm wide, oblong-linear, weakly compressed, 
dark brown, glabrous, with slightly raised ribs; pappus of numerous white, 
slightly barbed (under high magnification!) bristles, 7.5-8 mm long. 
Flowering (June)July to August (September). (Plate XXIII, Fig. 3). 

Montane forests, at 700-2,000(2,400) m. European Part: Upper 
Dniester (Carpathians); Caucasus: Ciscaucasia, Dagestan, Eastern 
Transcaucasia, Western Transcaucasia. General distribution^ Central 
Europe, Atlantic Europe, Mediterranean, Balkans-Asia Minor, Armenia 
and Kurdistan (Artvin District). Described from western Europe. Type in 
London. 

Note. The species Prenanthes tenuifolia L., described by Linnaeus 
from the Alps of southern Europe, has not been recognized as an 
independent taxon by the large majority of authors. However, the 
specimens from the Caucasus referred by us to P. purpurea differ slightly 
from the European members of this species by having more entire, narrow 
leaves, on the basis of which Ledebour referred the Caucasian plant to P. 
tenuifolia L. This difference is far from being consistent; therefore, the 
predominant point of view at present about the need for combining the 
two species into one appears to be correct. 

2. P. tatarinowii Maxim, in Mem. Acad. Petersb. div. sav. IX (1859) 
474 (Primit. Fl. Amur., sine numero, in nota); Kom. in Tr. Peterb. Bot. 
Sada XXV, 788 (Fl. Man'chzh. Ill, No. 1656); Kom. and Alis. Opred. 
Rast. Dal'nevost. Kr. II, 1102; Ling in Contrib. Inst. Bot. Nat. Acad. 
Peip. Ill, 4, 200; Kitam. in Mem. Coll. Sc. Univ. Kyoto, ser. B, XXIII, 1, 
154 (Compos. Jap. V). — Lactuca tatarinowii Franch. in Morot, Joum, de 
Bot. IX (1895) 293.— Nabalus tatarinowii Nakai in Fl. Sylv. Kor. XIV 
(1923) 116; Kitag. in Report. Inst. Res. Manch. Ill, App. 1, 460 (Lineam. 
fl. Manschur.). 

Perennial. Rhizome? Stem 70-110(160) cm high, paniculately 
branched above, 6-8 mm near base and here, usually with setose scaly 
hairs, more or less glabrous above to general inflorescence. Leaves thin, 
266 with distinct network of veins, on long petioles, as also lamina, more or 
less densely covered with scaly hairs or spinules; lower leaves lyrate, 
large, with hastate-cordate or triangular-hastate terminal segment, 
terminating in hastate cusp; segment unequally toothed with isolated 
spinules and scaly hairs in-between; lateral segments remote, one pair, 
ovate-oblong or oblong; middle cauline leaves smaller, lower or entire 
(without lateral segments), with less distinct cordate base; upper cauline 



265 

leaves irregularly rhombic or angular-ovate, uppermost leaves lanceolate 
or narrowly lanceolate. Capitula cylindrical, 9-10(12) mm long, on erect 
branches, in lax, often very large, paniculate general inflorescence, 
peduncles short, slender, densely covered with large or scaly glandular 
hairs, often with one to few reduced, scaly, reddish leaves. Involucre two- 
or three-rowed; outer involucral bracts very short, inner usually five, one- 
rowed, oblong-linear or linear, with isolated erect curly or scaly, but fine, 
usually glandular, hairs on dorsal side (under lens!). Florets in capitulum 
usually five; corolla light blue or pale with violet tips. Achenes about 4 
nmi long and 0.6-0.8 mm wide, terete, usually with five longitudinal, 
prominent ribs, dark brown, almost glabrous (at high magnification very 
short hairs visible), shghtly narrowed toward base, with broad apical disk 
bearing pappus of numerous thin, dirty-rusty or rusty, barbed, 6-7 mm- 
long bristles. Flowering July to September(October) (Plate XXIII, Fig. 5). 
In broad-leaved and mixed shady forests as well as on gravels along 
brooks and streams. — Far East: Ussuri (mostly southern part). General 
distribution: Japan?, China, Korean Peninsula. (Earlier reported from 
Japan, but, according to S. Kitamura, by mistake; besides, one doubtful 
sheet from Sakhalin Island is preserved in the Herbarium of the Botanical 
Institute, Academy of Sciences of the USSR.) Described from vicinity of 
Beijing. Type and isotype in Leningrad. 

3. P. cacaliifolia (M.B.) Beauverd in Bull. Soc. Bot. Geneve 2 Ser. 
II (1910) 114 and 115 ("f. cacaliaefolid"; seorsum impressum: Contrib. 
a I'etude des Compos.III, p. 16 and 17). — Sonchus cacaliaefolius M.B. 
Fl. taur.-cauc. II (1808) 241 and III (1819) 527. —Mulgedium 
cacaliaefolium DC. Prodr. VII, 1 (1838) 250; Ldb. Fl. Ross. II, 2 (1845- 
1846) 841 (incl. a. genuinum, (3. appendiculatum and y. giganteum Ldb.) 
Boiss. Fl or. Ill (1875) 801 (excl. minus Boiss.) — M. petiolatum C. Koch 
in Linnaea XVII (1843) 279. — Cicerbita cacaliaefolia Beuverd Grossh. 
267 Fl. Kavk. IV (1934) 253 and Opred Rast. Kavk. 514; Schchian in Fl. 
Gruzii VIII, 672; Askerova in Fl. Azerb. VIII, 545.— Exs.: Balansa., PI. 
d'Or. (Reliq. Mailleanae) No. 1466 (sub Mulgedio lazico Boiss.). 

Perennial. Rhizome about 1 cm wide, usually short, hard. Stem 50- 
100(150) cm high, usually solitary, slightly sulcate, glabrous or 
subglabrous below, paniculately branched above and more or less densely 
covered with fine, scaly, glandular hairs. Leaves thin, scaly or more or 
less so; lower leaves hastate-cordate or deltoid-hastate with deeply sinuate 
cordate-reniform base, aristate, shallow and unequally sinuate-toothed, 
teeth soft and short-cuspidate, lamina green above and gray or glaucous 
beneath, with long, gradually expanded semiamplexicaul sheathing petiole, 
sometimes divided and then with one or two or, less often, few usually 
oblong segments; in very rare cases, lamina of lower leaves divided into 



266 

few segments or lobes; upper leaves on broad semiamplexicaul petioles, 
higly variable (oblong to lanceolate and linear), with strongly divided 
and toothed lamina, or.more or less entire with cusp; uppermost leaves 
strongly reduced. Capitula narrow-campanulate, (10)12-14(15) mm long, 
at apices of slender peduncles of paniculate inflorescence, usually 
exceeding capitula and densely covered with fine, scaly, glandular hairs. 
Involucre usually two-rowed; involucral bracts oblong-linear, subobtuse 
or obtuse, on back more or less densely covered with hairs-like peduncle, 
but sometimes light or dark- violet; florets in capitulum about 15; corolla 
light blue. Achenes about 5 mm long, and 1 mm wide, oblong-cylindrical, 
slighdy flattened, usually with five prominent ribs, light brown, densely 
covered with very short, light colored, upward-directed hairs (under high 
magnification!); pappus of numerous dirty white or light rusty, slightly 
barbed (under a hand lens!) hairs, reaching 7-8 mm in length. Flowering 
July to October. (Plate XXIII, Fig. 6.) 

Middle mountain and subalpine zones up to 2,500 m; deciduous 
(predominantly beech), mixed and coniferous forests, forested ravines, 
less often in thickets of shrubs and tall grass; rarely in humid forests of 
lower mountain zone. — Caucasus: Ciscaucasia, Dagestan, Eastern and 
Western Transcaucasia, ?Southem Transcaucasia (needs confirmation!). 
General distribution: Balkans-Asia Minor (Lazistan, Trabzon). Described 
from Georgia. Type in Leningrad. 

Note. The species is higly variable in the shape and size of the leaves, 
as well as in the size of the plant as a whole. This provided grounds to 
Ledebour to distinguish three varieties, which, however, could hardly rank 
above a form and do not have serious taxonomic significance. 

268 4. P. pontica (Boiss.) Leskov in Tr. Bot. Muz. XXV (1932) 44.— 

Mulgedium ponticum Boiss in Ann. Sc. Nat. 4. Ser. II (1854) 248; id. in 
Tchihatch. As. Min. Botanique II (1860) 385. M. cacaliaefolium {3. minus 
Boiss. Fl. or. Ill (1875) 801. — Prenanthes cacaliaefolia Beauverd var. 
minor Beauverd in Bull. Soc. Bot. Geneve 2 Ser. II (1910) 115 (seorsum 
impressum: Contrib. a I'etude des Compos. Ill, p. 17). — Cicerbita pontica 
Grossh. Fl. Kavk IV (1934) 250 and Opred. Rast. Kavk. 514; Kolak Fl. 
Abkhazii IV, 300; Schchian in Fl. Gruzii VIII, 671. — P. cacaliaefolia 
Beauverd ssp. minus Stebbins in sched. (1938). — Ic: Tchihatch. As. Min. 
Atlas (1860), t. 34 (ic. optima!); Kolak, op. cit. Plate XXIX (ic, admodum 
mediocris). — Exs.: GRF No. 323 (sub Mulgedio cacaliaefolio). 

Perennial. Rhizome mostly short. Stem 30-50(70) cm high, slender 
(usually 1-3 mm thick), simple and weakly pubescent to glabrous below, 
somewhat paniculately branched above and densely covered with very 
fine, often violet, scaly, glandular hairs. Leaves thin, crowded mainly in 
lower part of stem, hastate-cordate, cordate-ovate or deltoid-hastate with 



267 

deeply emarginate base, acute, unevenly sinuate-toothed with teeth softly 
short-cuspidate; lamina green above, glaucous beneath, with flat, 
semiamplexicaul, slightly sheathing, petiole at base almost equaling lamina 
or longer; upper leaves usually one to few, sessile, often comprising 
expanded petiolar base and short, reduced, divided lamina, or lamina more 
or less entire, oblong-lanceolate, linear-lanceolate or linear; uppermost 
leaves (on inflorescence branches and peduncles) scaly. Capitula cylindrical 
or narrow-campanulate, about 12-13 mm long, at apices of slender, usually 
longer, peduncles of well-developed inflorescence than capitula and 
densely covered with fine scaly glandular hairs. Involucre two- or three- 
rowed; involucral bracts often violet, outer bracts on back covered with 
hairs like peduncles; inner bracts longer, obtuse, with fewer glandular 
hairs, or eglandular. Florets in capitulum usually 8-10; corolla light blue. 
Achenes about 5 nmi long and 1 mm wide, oblong-cylindrical, slightly 
flattened, usually with five distinctly raised longitudinal ribs and large 
number of inconspicuous or almost indistinct ribs in-between, light brown, 
densely covered with very short, light colored, upward-directed hairs 
(under high magnification!); pappus of numerous dirty white or hght rusty 
barbed (under a lens!) bristles, 7-8 mm long. Flowering August to October. 
(Plate XXIII, Fig. 7.) 

Mountain forests and edges up to subalpine zone. — Caucasus: 
Ciscaucasia (Ossetia), Western Transcaucasia (Black Sea Coast — Sochi 
269 District, Abkhazia, Colchis, Kutaisi District, Adzharia), ?Eastern 
Transcaucasia (Zakataly District; imprecise data). General distribution: 
Balkans-Asia Minor (Pontic Range). Described from Turkey. Type in 
Geneva. 

Note. A species very closely related to P. cacaliifolia with which, 
possibly, it hybridizes in areas of contact. However, P. pontica differs 
from P. cacaliifolia by having a smaller size, more slender stem, lamina 
of the lower leaves reaching the base and usually (3)5-8(10) cm long (in 
P. cacaliifolia, lamina, as a rule, larger); the petiole usually only slightly 
expanded at the base; leaves mainly crowded in the lower part of stem; a 
smaller number of florets in the capitulum; a comparatively weakly 
developed inflorescence and several other features. These morphological 
differences and the distinct geographic range compel us to consider P. 
pontica as a separate species and not include it as a subordinate taxon 
under P. cacaliifolia. 

Subgenus 2. Nabalus (Cass.) Babcock, Stebbins and Jenkins in 
Cytologia, Fujii Jubil. vol. (1937) 190 — Nabalus Cass, in Diet. sc. nat. 
XXXIV (1825) 94.— Prenanthes sect. Nabalus Kitam. in Mem. coll. Sc. 
Univ. Kyoto, ser. B, XXIII, 1 (1956) 152, p. p. (Compos. Jap. V).— 
Capitula with 25-35 flowers; corolla yellow or whitish; achenes yellow 



268 

or dark brown, terete with numerous longitudinal ribs. Primarily a New 
World plant. 

Lectotype of subgenus: Prenanthes alba L. 

5. P. maximowiczii Kirp. nom. nov. — Nabalus ochroleucus Maxim, 
in Bull. Acad. Sc. Petersb. XV (1870) 376. — Prenanthes ocroleuca Hemsl. 
in Joum. Linn Soc. XXIII (1888) 486 (non Rafm., 1824); Kom. in Tr. 
Peterb. Bot. Sada XXV, 789 (Fl. Man'chzh. Ill, No'. 1657); Kom. and 
Alis. Opred. Rast. Dal'nevost. Kr. II, 1102; Kitam. in Mem. Coll. Sc. 
Univ. Kyoto, ser. B, XXIII, 1, 153 (Compos. Jap.). — Lactuca ochroleuca 
Franch. in Morot, Joum. de Bot. IX (1895) 293. 

Perennial. Plants often with strong rhizome. Stem 70-100(120) cm 
high, sulcate, up to 9 mm thick at base, in lower part densely covered 
with scaly, dark brown, hairs or (less often) more or less glabrous, 
branched above and covered with crisped, less often mixed with scaly 
setose, hairs. Lower leaves (early- withering) and middle cauline leaves 
large, lyrate-pinnatisect, on very long, winged, expanded at base and 
semiamplexicaul petiole, usually covered beneath with scaly hairs; terminal 
segment triangular or irregularly rhomboid, lateral segments two to four, 
oblong or broadly ovate, or rhomboic, sometimes irregular in shape; all 
segments acute, irregularly coarsely toothed-sinuate, with teeth terminating 
270 into obtuse spinules; upper leaves strongly reduced, sessile, usually 
lanceolate or oblong, often densely setose at base. Capitula with 25-30 
florets, narrow-campanulate, at fruiting about 15 mm long, on unequal 
peduncles, densely covered with short crisped hairs (under a lens!) and 
often in compact racemose-paniculate inflorescence. Involucre two- or 
three-rowed; involucral bracts dirty green, with scaly erect hairs on back. 
Corolla yellow. Achenes 7-8 mm long and about 0.6 wide, terete, with 
numerous, prominent longitudinal ribs, yellow or dark brown, very densely 
covered with light colored, upward-directed, short hairs (under high 
magnification!), somewhat distinctly narrowed toward base, with almost 
quadrate, smooth, corona-like process at apex, bearing pappus of 
numerous, dirty-rusty or rusty, finely barbed 6-7(8) mm-long bristles. 
Flowering August to September (Plate XXIII, Fig. 9). 

Wet meadows, along streams and near springs, moist forest glades, 
and also shrub thickets on marshy soil. — Far East: Ussuri (southern part). 
General distribution: Japan and China (northeastern China), Korean 
Peninsula. Described from vicinity of Vladivostok. Type and isotype in 
Leningrad. 

Note. It has always been said that the first date of publication of the 
basionym of this species was in 1871. However, according to the data on 
page 384 of Vol. XV, ""Bull Acad. Sc. Petersb., " this publication came 
off the press on November 11, 1870. 



269 




271 Plate XVH. 

1 — Lactuca wilhelmsiana Fisch. and Mey., habit and achene; 2 — L. raddeana Maxim., 
achene; 3 — L. striata Waldst. and Kit., achene; 4 — L. indica L., achene; 5 — L. triangulata 

Maxim., achene. 



270 

Species of Doubtful Taxonomic Position 

6. P. abietina (Boiss.) Kirp. comb. nova. — Mulgedium abietinum 
Boiss. and Bal. ex Boiss. Fl. or. Ill (1875) 802. — Cicerbita abietina 
Beauverd ex Grossh. Fl. Kavk. IV (1934) 252 and Opred. Rast. Kavk. 
514.— C. abietina Stebbins in Joum. Bot. Lond. LXXV (1937) 16.— 
Lactuca abietina Bornm. in Mitt. Thur. Bot. Ver. n. s. XX (1904-1905) 
25.—Crepis abietina Beauverd in Bull. Soc. Bot. 2 Ser. II (1910) 115; 
Kolak. Fl. Abkhazii IV, 308; Schchian in Fl. Gruzii VIII, 700. 

Perennial. Rhizome about 1 cm thick, annually producing solitary 
herbaceous stem from apex; previous year's stems preserved as fragments 
with remnants of leaves; adventitious roots slender, cord-like to almost 
272 filiform, fewer. Stem 90-150 cm high, 5-10 mm thick at base, weakly 
branched near apex, sulcate, fistular, entirely or almost entirely glabrous. 
Leaves petiolate; lower leaves on long petioles as long as or exceeding 
lamina, middle leaves on shorter petioles almost half as long as lamina, 
and upper leaves short-petiolate or more or less sessile; lamina very thin, 
almost chartaceous, with distinct network of veins, rotund-triangular or 
triangular, more or less acuminate, cordate-hastate at base, margin broadly 
but shallow sinulate or coarsely large and obtusely toothed, with thin soft 
spinose processes; less often petioles with small leafy auricles. Capitula 
ovoid or campanulate, 5-10 mm wide and about 1 cm long, few, on long 
peduncles slightly expanded above (below capitula), in lax paniculate- 
corymbose inflorescence; sometimes capitula arising from axils of upper 
leaves, less often only few or solitary. Involucral bracts two- to four- 
rowed linear-lanceolate, dirty dark green to almost black. Capitula with 
30-35 florets; corolla light- or bright-yellow, two times or more as 
long as capitulum. Achenes about 4 mm long and 1 nrni wide, slightly 
flattened, almost terete or indistinctly four- or five-angled, slightly 
narrowed toward apex and near base, usually with four or five more 
strongly protecting ribs and indefinite number of less conspicuous ribs, 
dark brown, densely covered with very short, light-colored hairs (under 
high magnification!); pappus of snow-white, almost entirely smooth, 
persistent hairs, slightly exceeding achene. Flowering (June)July to August. 
(Plate XXIII, Fig. 4.) 

Montane forests to their upper limit, less often in tall-grass and 
secondary mountain meadows; mostly under forest cover (beech, fir, pine; 
elfin woodland) or in thickets of rhododendron. — Caucasus: Ciscaucasia, 
Eastern Transcaucasia, ?Westem Transcaucasia. General distribution: 
Balkans-Asia Minor (Asia Minor). Described from Lazistan. Type in 
Geneva. 

Note. Many authors have included this species under the genus 
Cicerbita, from which it differs by the absence of an outer pappus, the 



271 

firmly attached hairs of the inner pappus, a larger number of florets in 
capitulum, and other characters. 

According to Boissier (op. cit. 803), this plant occupies an 
"intermediate position between Crepis and Mulgedium, " but "because of 
the appressed achenes, which are smaller than the persistent pappus," it 
should be placed in the genus Mulgedium (despite the persistent pappus). 
Actually the achenes are little or, often, not at all compressed. 
Morphologically, this species is similar to some species of Crepis, but 
the possibility of its belonging to this genus was refuted by such 
distinguished specialists of Crepis as Babcock and Stebbins. 

At present, it is most appropriate to assign this species to the genus 
Prenanthes. to which it is closest in the structure of its achenes. True, the 
long overdue revision of Prenanthes, whose heterogeneous nature was 
273 reported by botanists already at the beginning of the 19th century, may 
introduce significant changes even in the position of the species under 
consideration. 

7. P. angustifolia Boulos in Bot. Notis. CXV (1962) 59.— Sonchus 
dentatus. Ldb. Ic. pi. fl. Ross. 1 (1829) 21; Ej. Fl. Alt. IV (1833) 141; 
DC. Prodr. VII, 1, 187; Ldb. Fl. Ross. II, 2, 835; Kryl. Fl. Zap. Sib. XI, 
3019; Sobolevskaya, Konsp. Fl. Tuvy 200; Grubov, Konsp. Fl. MNR 
282— No. 1854 (Tr. Mong. Kom. Vyp. 67).— Ic: Ldb., op. cit. (1829) t. 
87 (color., bona). 

Perennial. Rhizome more or less long, 2-6(10) mm thick. Stem (7)10- 
50(65) cm high, slender, indistinctly sulcate, glabrous or subglabrous, 
more or less strongly (sometimes right from base) branched. Leaves 
crowded in lower part of stem, oblong-elliptical, linear-lanceolate or linear, 
shallow sinuate-toothed or toothed, sparsely spinulose, on short, narrow- 
winged, semiamplexicaul petiole; middle and upper leaves distant, entire, 
sessile, strongly reduced to scaly, with fertile axillary branches. Capitula 
with about 15 florets, usually less numerous, in corymbose or corymbose- 
paniculate inflorescence; peduncles unequal, slender, covered with short 
crisped or longer scaly hairs, or glabrous. Involucre oblong-cylindrical or 
narrow-campanulate, about 10 mm long, 4-7(10) mm wide, usually two- 
rowed; involucral; bracts dark, dirty green, linear or oblong-lanceolate, 
on back covered with long, narrow, scaly, yellowish-green, erect hairs or 
glabrous; inner bracts with barbules of thin light colored hairs (under a 
lens!) at apex. Achenes oblong or oblong-ellipsoid, slightly flattened, dark 
violet, 4.5-6 mm long, 1-1.25 mm wide, with about 10 prominent 
longitudinal ribs, and light yellowish constriction at apex, covered with 
numerous, very short, upward-directed hairs (under a hand lens!); pappus 
of fine, white or shghtly grayish, weakly barbed (under a lens!) brisdes, as 
long as achene or longer. Flowering June to August. (Plate XXIII, Fig. 8.) 



272 

On more or less wet saline soils; meadows, banks of reservoirs — 
Western Siberia: Altai; Eastern Siberia: Angara-Sayan (Tuva ASSR)? 
General distribution: Mongolia. Described from Altai (Chuya River). Type 
in Leningrad. 

Note. Considering the achene structure, number of florets per 
capitulum, nature of the Involucral bracts and other characters, this species 
cannot be^kept in the genus Sonchus. It occupies an intermediate position 
between Prenanthes and subgenus Mulgedium of Lactuca, but is somewhat 
closer to the former. P. angustifolia differs from typical Prenanthes by 
the yellow color of the corolla and the leaf shape; and from Mulgedium, 
again by the corolla color (in Mulgedium it is light blue), as well as the 
achene shape. 
274 The name P. dentata cannot be used in view of the existence of an 

earlier homonym. 

The plants vary markedly in the nature of the pubescence, extent of 
branching, and growth vigor. A form with involucral bracts lacking scaly 
hairs is found in the Tuva ASSR and in Mongolia. Most such plants 
were labeled Sonchus dentatus Ldb. var. glabrisquameus Krasch. and Ujin" 
by Iljin in the Herbarium of the Botanical Institute, Academy of Sciences 
of the USSR. 

Doubtful and Deleted Species 

1. P. alata (Hook.) Dietr. Synops. plant. IV (1847) 1309.— Nabalus 
alatus Hook. Fl. Bor. Amer. I (1833) 294, t. 102; Ldb. Fl. Ross. II, 2, 
840. — Sonchus hastatus Less, in Linnaea, VI (1831) 99. — Mulgedium 
hastatum Less. Synops. Compos. (1832) 142. 

This species should be excluded from the Flora of the USSR. For 
details, see V.L. Komorov, Fl. P-va Kamchatki [Flora of Kamchatka 
Peninsula] III, 1930, p. 207, and in E. Hulten [Fl. of Kamtchatka and the 
adjacent Islands IV (1930) 232]. 



GENUS 1654. Lactuca L.'-^ 

L. Sp. pi. (1753) 795; DC. Prodr. VII, 1 (1838) 133; Benth. in Benth. 
and Hook. f. Gen. pi. II (1873) 524, p. p.; O. Hoffm. in Pflanzenfam. IV, 

5 (1894) 371, p. p. 



' Treatment by M.E. Kirpicznikov. 

^ From the Latin word lac — milk; the name indicates the presence of milky latex in 
the plants. 



273 

Capitula homogamous, with (9)10-20(27) florets, small (up to 
(5)9-14(20) mm long at fruiting), usually narrow-cylindrical, oblong, 
truncate-turbinate or campanulate, sometimes constricted near middle or 
base, in paniculate, corymbose- or racemose-paniculate or, less often, 
spicate inflorescence. Involucral bracts herbaceous, imbricate, three- or 
four-rowed; inner bracts several times as long as outer ones, usually 
scarious along border. Receptacle smooth. Corolla yellow, less often light 
blue, blue, lilac or pink. Achenes more or less flattened to flat, narrow- 
obovoid, oblong-ellipsoid, oblong, oblong-linear, oblong-ellipsoid or 
ellipsoid, differendy colored (brown of various shades, dark violet, blackish 
and other shades), with one to many or somewhat numerous prominent 
longitudinal ribs, often slightly rugose due to short hairs in transversely 
undulate rows (under a hand lens!), terminating into long filiform beak, 
275 differing in color and structure from rest of achene, or upper part narrowed 
in thin beak-like structure, colored like rest of achene; less often achene 
apex narrowed into somewhat thick crown or not; pappus bearing disk 
wide, usually equaling broadest part of achene or slightly broader; pappus 
of white, less often somewhat dirty or rusty thin hairs, usually falling 
singly, less often more or less firmly attached to disk. 

Annual, biennial and perennial herbs, less often semi-shrubs, 
inhabiting the forest zone and mountains, as well as more or less arid 
steppes, semideserts and deserts on soils with various degree of salinity. 

Lectotype of genus: Lactuca sativa L. 

The genus includes 100-150 species, a large number of them 
distributed in Asia; many occur in the Mediterranean countries and tropical 
Africa; relatively few species are found in North America, and only a 
few have been reported from Central America. 

Note. For the sake of practical convenience, we are adopting a quite 
broad concept of the genus Lactuca. In the context of the present approach 
to phylogeny, it probably would be more appropriate to consider the four 
subgenera as separate genera. 

1. Rhizome slender, creeping, terminating in slender and short 
vanishing stem up to 8-15 cm high including leaves; leaves two 
to few, on more or less long (usually about 5 cm) petioles; capitula 
in ones and twos, less often a few, on axillary peduncles. Plants 
endemic to Talass Alatau 20. L. mira Pavl. 

+ Plants without creeping rhizome, or leaves many, and capitula not 
axillary 2. 

2. Achenes flat, ellipsoid or oblong-ellipsoid, 2-2.5 mm wide in 
broadest part, less often 1 .5 mm, almost black or dark violet; aerial 
parts of plant with narrow-fusiform tuberous thickenings. Within 



274 

the USSR, plants growing only in the Far East, less often also in 

Eastern Siberia 3. 

+ Achenes usually flattened, up to 1 mm wide, less often to 1.5 
mm, differently colored; aerial parts of plant without narrow- 
fusiform tuberous thickenings 5. 

3. Achenes with very short but distinct beak; capitula with 20-25 
florets, 13-15 mm long at fruiting 7. L. indica L. 

+ Achenes with very short, somewhat broad neck, but without distinct 
beak; capitula with 10-15 florets, 10 mm long at fruiting 4. 

4. Achenes with few prominent longitudinal ribs on each side; leaves 
276 on both sides and along margin (under a lens!) with short 

squamiform hairs, most distinct along veins beneath, unevenly 
spinose-toothed along margin and usually with narrow .reddish 
frill; stem rather densely covered with narrow squamiform hairs 
up to point of forking 9. L. raddeana Maxim. 

+ Achenes with single, rarely two prominent longitudinal ribs on 
each side; leaves usually with dense, narrow, pale, squamiform 
hairs (under a hand lens!) sometimes above and along unevenly 

spinose-toothed margin 8. L. triangulata Maxim. 

5(2) Beak or narrow beak-like part more or less as long as achene or 
longer, usually filiform and almost always differently colored than 
body of achene 6. 

+ Beak (or narrowed beak-like part) half as long as achene or slightly 
longer, usually concolorous with body of achene; sometimes 
achenes without beak 15. 

6. Beak colored like whole achene. Plants rather large (50-130 cm 
high), with undivided, thick (almost coriaceous), elliptical or 
oblong-obovate leaves, punctate beneath. Plants growing in 
Badkhyz and Kugitang 19, L. spinidens Nevski 

+ Beak differently colored than achene. or more or less colorless. 
Plants smaller or leaves different 7. 

7. Base of beak with two pendent rod-shaped appendages 
(deliquescent in water!) at achene apex; as a result beak appears 
to lie on top of achene; beak three to four times as long as achene. 
Annual, distributed all over Soviet Central Asia, as well as southern 
Transcaucasia and Western Siberia 18. L. undulata Ldb. 

+ Base of beak without rod-shaped appendages; beak arising from 
achene apex or upper neck-like part of achene 8. 

8. Achene terminating in roundish notch where conically thickened 
base of beak originates, narrowed above into filiform structure; 
achenes usually with three longitudinal ribs on each side. Annual. 
Plants of Soviet Central Asia (almost exclusively in Tien Shan 
and Pamiro-Alai districts) 17. L. auriculata DC. 



275 

+ Achene not terminating in roundish notch; base of beak without 
conical thickening; achenes with one or many (five-seven-nine or 
more) longitudinal ribs on each side 9. 

277 9. Short annual plants, (2)7-15(20) cm high; basal leaves in rosette, 

with rather long, crisped or flexuous, light colored hairs beneath; 
achenes light colored (yellowish or dark brownish-yellow) with 
single prominent rib in middle on each side. Plants of Soviet 

Central Asia 16. L. glauciifolia Boiss. 

+ Plants taller, or leaves without crisped hairs beneath, and achenes 
of different color 10. 

10. Achenes with single prominent rib in middle on ventral and dorsal 
sides, dark or blackish-brown, almost entirely glabrous (rather short 
hairs visible only under high magnification). Short (10-25 cm 
high), biennial or perennial plants, rarely found (within the USSR, 
so far known only from the western part of the Kara-Kum District) 

15. L. rosularis Boiss. 

-I- Achenes with more or less numerous prominent longitudinal ribs 
on each side 11. 

11. Achenes about 1.5 mm wide (in broadest part), dark violet to 
almost black, abruptly narrowed at apex into about 1 mm-long 
narrow part (neck) bearing fiUform beak almost as long as achene; 
pappus readily detached. Stem up to 7-10 mm thick at base, fistular 
14 L. georgica Grossh. 

-I- Achenes narrower, or their beak longer 12. 

12. Achenes, including the upper part, covered with fine, very short, 
upward-directed, stiff hairs (under a hand lens!); capitula borne 
singly on slender, virgate branches in lax spicate racemes; 
peduncles slender, short or almost undeveloped, and then capitula 
subsessile or sessile; beak usually one and one-half to two times 
as long as achene. Plants from the European part of the USSR 
and Caucasus 12. L. saligna L. 

-1- Achenes in upper part covered with more or less soft, often squar- 
rose, transparent hairs (under a lens!) 13. 

13. Lower leaves undivided, rotund-obovate or elliptical, sometimes 
sinuate; achenes usually gray (less often dark or light brown), about 
4 mm long and 0.8-1 mm wide; pappus almost as long as achene 
and more or less firmly attached. A universally cultivated plant, 
but sometimes growing in the wild ILL. sativa L. 

+ Lower leaves variously divided (runcinately pinnatisect, 
pinnatipartite, pinnatifid, erose-pinnatipartite or pinnatilobate), if 
entire, then lanceolate 14. 

278 14. Lower and middle cauline leaves usually runcinate, completely 

surrounding stem (in clear weather held vertically in sunny places); 



276 

peduncles usually longer than capitula; beak slightly longer or one 
and one-half to two times as long as achene; pappus about 6 mm 
long, of not always deciduous hairs. Plants widely distributed in 
the European part of the USSR, all districts of Caucasus and Soviet 

Central Asia, as also in Siberia 10. L. serriola Tomer 

+ Lower leaves usually runcinate, or lower and middle cauline leaves 
lanceolate and lanceolate-Unear to linear and (very rarely) oblong; 
peduncles usually short, as a result, capitula often subsessile or 
even sessile; beak more or less as long as achene, sometimes 
distinctly longer; pappus about 5 mm long, of rather readily 
detached hairs. Plants of southern part of Volga Region, Caucasus, 

Western Siberia, and Soviet Central Asia 

13. L. altaica Fisch. and Mey. 

15(5) Corolla yellow; achenes dark violet to almost black with 
concolorous upper beak-like part, half (or slightly more) as long 

as achene 18. 

+ Corolla blue or light blue (but color of Lactuca winkleri unknown); 
achenes usually light colored (yellowish-brown, olive colored, etc.), 
less often blackish 16. 

16. Achenes yellow or sand-colored, with dark brown annulus at apex, 
densely covered with short, light-colored hairs throughout (under 
a lens!); pappus hairs very readily detached. Plants endemic to 
Pamiro-Alai 3. L. winkleri Kirp. 

+ Achenes dark, with indistinct annulus, wholly covered with short 
hairs, usually arranged in transverse-sinuate rows (under a lens!); 
pappus hairs more or less readily detached 17. 

17. Longitudinal ribs in middle; as a result, relatively thick margins 
of achenes stand out; beak-like narrowing either not developed, 
or one-fourth as long as achene (very rarely longer); cauline leaves 
usually with broad amplexicaul base, thin 

1. L. sibirica (L.) Benth. ex Maxim. 

+ Longitudinal ribs more or less uniformly distributed on achene; 
beak-like narrowing often (especially in upper part) lighter-colored, 
usually one-third to half as long as achene; cauline leaves often 

narrowed toward base, usually stiff and thickish 

2. L. tatarica (L.) C.A. Mey. 

18(15) Middle cauline leaves entire or subentire 

5. L. chaixii Vill. 

279 + Middle cauline leaves pinnatipartite or pinnatisect, 19. 

19. Achenes with five or more prominent longitudinal ribs on each 
side, gradually narrowed toward apex into about 3 mm-long beak; 
segments of leaves usually narrow-triangular or linear-lanceolate; 
roots usually tuberous 6. L. wilhelmsii Fisch. and Mey. 



277 

+ Achenes with two or three, less often more (but usually not more 
than five), prominent longitudinal ribs on each side; narrowed 
toward apex into about 2 mm-long beak; segments or lobes of 
leaves mostly oblong or ovate-lanceolate; roots usually not 
tuberous 4. L. stricta Waldst. and Kit. 

Subgenus 1. Mulgedium (Cass.) Babcock, Stebbins and Jenkins in 
Cytologia, Fujii Jubil. vol. (1937) 190 p. min. p. — Mulgedium Cass, in 
Diet. sc. nat. XXXIII (1824) 296 and XLVIII (1827) 426, p. p.— 
Agathyrsus D. Don in Edinb. New Phil. Joum. VI (1829) 310 p. p.— 
Mulgedium sect. 1. Eumulgedium DC. Prodr. VII, 1 (1838) 248, p. min. 
p. — Lactuca sect. 4. Cicerbita Benth. in Benth. and Hook. f. Gen. pi. II 
(1873) 525, p. min. p. — Mulgedium sect. I. Lactucopsis (Sch. Bip.) Boiss. 
Fl. or. Ill (1875) 799. — Lactuca sect. Mulgedium Clarke, Compos. Ind. 
(1876) 260, p. p. — Cicerbita § 1. Mulgedium Beauverd in Bull. Soc. 
Geneve, 2 Ser. II (1910) 116 (seorsum impressum: Contrib. a. I'etude des 
Compos. Ill, p. 18). — Lactuca sect. Lactucopsis Kitam. in Acta Phytotax. 
et Geobot. XI, 2 (1942) 125. — Lagedium Sojak in Novit. bot. Horti bot. 
Univ. Car. Prag. (1961) 34. — Achenes slightly flattened, yellowish-brown 
to dark olive and black, with few or more or less numerous prominent 
longitudinal ribs on each side, rather densely covered with very short, 
stiff, upward-directed hairs (under a hand lens!); achene apex attenuate 
into more or less distinct neck (sometimes rather thin and then looking 
like beak), concolorous with achene, or (less often) slightly hghter-colored; 
collar usually one-eighth to one-fourth as long as achene, less often up to 
half and more but, in some cases, neck almost not developed, and then 
achene apex directly modified into pappus-bearing disk; pappus of dirty 
white or dirty yellow, less often rusty or pure white hairs, falling 
individually; corolla light blue, blue, or lilac. Perennial, mostly weeds or 
semi-weeds, usually with strongly developed root system and rhizomes. 

Type of subgenus: Lactuca tatarica (L.) C.A. Mey. 
280 Note. This characterization of the subgenus is quite tentative, as it is 

based on the analysis of characters of only the local members of this 
taxon (the scope of the subgenus Mulgedium as a whole is not yet definite). 

1. L. sibirica (L.) Benbh. ex Maxim, in Bull. Acad. Sc. Petersb. 
XIX (1874) 528; Schmalh. PI. II, 166; Kitag. in Report Inst. Sc. Res. 
Manch. Ill, App. 1, 456 (Lineam. fl. Manshur.); Sugaw. 111. fl. Saghal. 
IV, 1927; Kryl. Fl. Zap. Sib. XI (1949) 3024 (incl. var. integrifolia (Ldb.) 
Serg. and var. dentata (Ldb.) Serg. op. cit. p. 3025); Sobolevskaya, Konsp. 
Fl. Tuvy 201; Grubov. Konsp. Fl. MNR 282, No. 1857 (Tr. Mong. Kom. 
Vysh. 67); Kitam. in Mem. Coll. Sc. Univ. Kyoto, ser. B, XXIII, 1, 134 
(Compos. Jap. V); Karav. Konsp. Fl. Yakutii 178. — Sonchus sibiricus L. 



278 

Sp. pi. (1753) 795; Ldb. Fl. Alt. IV (1833) 140 (incl. a. integrifolius 
Ldb. and p. dentatus Ldb. op cit. p. 141). — Mulgedium runcinatum Cass. 
in Diet. sc. nat. XXXIII (1824) 296, p. p. — Agathyrsus sibiricus D. Don 
in Edinb. New Phil. Joum. VI (1829) 310. — Mulgedium sibiricum Cass, 
ex Less. Synops. Compos. (1832) 142; DC. Prodr. VII, 1 (1838) 249 
(excl. p. indivisum DC. ibid); Ldb. Fl. Ross. II, 2 (1845-1846) 844 (incl. 
a. integrifolium Ldb. and (3. dentatum Ldb.); Turcz. in Bull. Soc. Nat. 
Mosc. XXI, 3, 124 (Fl. baic.-dahur. II, No. 721); Maxim, in Mem. Acad. 
Petersb. div. sav. IX, 182 (Primit. Fl. Amur. No. 466); Rgl. in Mem. 
Acad. Petersb. ser. VII, 4, 98 (Tentam. Fl. Ussur. No 4. 311, errore sub 
Mulgedio sibirico L.); Kom. in Tr. Peterb. Bot. Sada XXV, 773 (Fl. 
Man'chzh. Ill, No. 1643); Fedtsch. and Fler. Fl. Evrop. Ross. 1055; O. 
and B. Fedtsch. Perech. Rast Turk. IV, 351 (Consp. Fl. Turk. No. 2859); 
Lindem. Svensk fanerogam. fl. Andra uppl. 559; Kom. Fl. P-va Kamchatki 
III, 187; Hult. in Kungl. Sv. Vet. Akad. Handl. Ser. 3, Bd. 8, 2, 229 (Fl. 
of Kamtch. IV, No. 777); Kom. and Alis. Opred. Rast. Dal'nevost. Kr. 
II, 1097; Hiit. Suom. Kasv. 728; Kraschen. in Fl. Yugo-Vost. VI, 464; 
Perf. Fl. Sev. Kraya II-III, 374; Leskov, Fl. Malozem. Tundry, 103 (Tr. 
Sev. Bazy Akad. Nauk SSSR, Vyp. 2); Govorukhln, Fl. Urala 521; Kirp. 
in Majevski, Fl. (1954) 630; Vorobev in Tr. Dal'nevost. Fil. Ser. Bot. Ill 
(V) 77 (Materialy k Fl. Kuril'skikh o-vov. No. 976); Stank, and Tal. 
Opred. Vyssh. Rast. (1957) 442; Popov, Fl. Sr. Sib. II, IKl .—Mulgedium 
kamtschaticum Ldb. in Denkschr. Konigl.-Bayer. Bot. Gesellsch. Regensb. 
Ill (1841) 65 (Commentarius in J.G. Gmelini Floram sibiricam); id. Fl. 
Ross. II, 2, 845. — Sonchus maritimus Pall, ex Ldb. Fl. Ross. II, 2 (1845- 
1846) 843. — Lagedium sibirium Sojak in Novit. bot. Horti bot. Univ. Car. 
Prag. (1961) 34.— Ic: J.G. Gmel. Fl. Sibir. II (1749), t. Ill, fig. A and B; 
Schkuhr, Bot. Handb. Ill (1803), t. CCXVII (color., mediocr.); Sugaw. 1. 
c. t. 887.— Exs.: Herb. Fl. Ingr. No. 366; GRF No. 1229 (utroque casu 
sub Mulgedio sibirico). 
281 Perennial. Rhizomes mostly vertical, sometimes branched. Stem 

(10)30-70(110) cm high, usually solitary, almost smooth or weakly sulcate 
to ribbed, glabrous, often reddish, rather densely and uniformly leafy (fully 
developed leaves reaching, at least, base of common inflorescence), more 
or less branched in upper part. Leaves lanceolate or oblong-lanceolate, 
acute, sessile; lower leaves basally narrowed, others with semiamplexicaul 
or almost amplexicaul, cordate, cordate-auriculate or sagittate base; lamina 
green above, glaucous-green beneath, entire or subentire, with narrow 
bent down frill (var. integrifolia (Ldb.) Serg.), or from weakly emarginate- 
toothed to more or less strongly pinnatifid or even runcinate (var. dentata 
(Ldb.) Serg.), lobes (parts or segments) often falcate. Capitula with about 
20 florets, globose turbinate or campanulate, 9-13 mm long with fruits, 
in lax corymbose or corymbose-paniculate inflorescence; peduncles 



279 

slender, glabrous, usually longer than capitula. Involucral bracts three- or 
(less often) four-rowed, often purple, lilac or mottled (with dark spots), 
with very fine, dense, appressed pubescence on dorsal surface (under a 
hand lens!), inner bracts subobtuse, often with frill of thin, light-colored 
hairs. Corolla blue or light blue (very rarely white — albino). Achenes 4- 
5 mm long and 1-1.25 mm wide, oblong-linear or elliptical, somewhat 
flat, yellowish-brown to dark olive or blackish, with few, prominent, 
longitudinal ribs in middle or achene, rather densely covered with very 
short, light-colored, stiff hairs (under high magnification!), and, sometimes, 
also with few, rather long, white hairs at base; achenes narrowed above 
into more or less distinct neck, usually, not exceeding one-fourth of achene 
length and terminating in funnel-shaped broadening with roundish or four- 
or five- obtusely-angled disk; pappus of numerous, thin, dirty white or 
rusty (in rare cases pure white) hairs, weakly but quite frequently toothed 
(under a hand lens!), 5-7 mm long, slightly connate at base or free, in 
both cases easily detached from disk. Flowering June to July. (Plate XVI, 
Fig. 3.) 

Semiweeds, widely distributed in forest region, found in forest and 
floodplain meadows, thickets of willows and shrubs along forest edges 
on logged areas and burns, in forests (especially open forests), near 
irrigation channels and marshes (less often in marshes), as well as along 
old fields and wastelands and sometimes in cultivated fields; much less 
often found in steppe region, mostly near forest islands, on meadows, 
sometimes in thickets of wormwood or other nonforested herb or herb- 
shrub communities. — Arctic: Arctic Europe, Chukotka! (rarely!); European 
Part: Karelia-Lapland (south), Dvina-Pechora, Baltic Region (very rarely), 
Ladoga-Ilmen, Upper Volga (rarely!), Volga-Kama, ?Middle Dnieper (one 
sheet from vicinity of Ternopol), Trans- Volga Region (rarely!); Western 
Siberia: all regions; Eastern Siberia^ all regions; Far East: all regions; 
Soviet Central Asia: ?Dzhungaria-Tarbagtai (one doubtful sheet from Kopal 
Mountain). General distribution: Scandinavia, Mongolia, Japan and 
282 China, ?North America (continually reported earlier; not seen in herbarium, 
and did not find precise information in more recent publications). — 
Described from Siberia and Scandinavia ("Sibiria, Suecia, Finlandia"). 
Type in London. 

Note. A species that varies relatively little, apparently with a few 
ecological races. To date there is almost no precise information available 
on the morphology of the underground parts of the plants. 

Komarov (Fl. Poluostrova Kamchatki [Flora of Kamchatka Peninsula] 
III, 1930, p. 187) included "Youngia sibirica Erman Verz. 48" among 
the synonyms of Mulgedium sibiricum. I did not find this name in the 
publications of Erman, and it is also not mentioned in Index Kewensis, or 
in the work of Hulten on the Flora of Kamchatka. Erman listed only 



280 

Sonchus sibiricus L. (cf.: Erman, Reise, Naturhist. Atlas (Verzeichn. etc.) 
1835, p. 57). 

Economic Importance. It has some fodder value (cf. Larin et al. 
Kormovye Rast. Senokosov i Pastbishch SSSR [Fodder Plants of the 
Grasslands and Pastures of the USSR], Vol. Ill, Sel'khozgiz, Moscow- 
Leningrad, 1956, p. 677). 

2. L. tatarica (L.) C.A. Mey. Verzeichn. Pfl. Cauc. (1831) 56, 
Schmalh. Fl. II, 166; Grossh. Fl. Kavk. IV (1934) 255 and Opred. Rast. 
Kavk. 515; Ling in Contrib. Inst. Bot. Nat. Acad. Peip. Ill, 4, 186; Kryl. 
Fl. Zap. Sib. XI (1949) 3023 (incl. var. glabra, and v2C[.floccosa Serg.); 
Schchian in Fl. Gruzil VIII, 683; Sobolevskaya, Konsp. Fl. Tuvy 201; 
Hayek in Hegi, 111. Fl. 2 Aufl. VI, 2, 1130; Grubov. Konsp. Fl. MNR 
283, No. 1858 (Tr. Mong. Kom Vyp. 67); Kitam. in Res. Kyoto Univ. 
Exp. Karak. a. Hinduk. 1955, II, 431 (Fl. of Afghanistan, 1960); Vveden- 
sky. et al. Opred. Rast. Golodnoi Stepi (40 Tr. Tashk. Gos. Univ;. Vyp. 
178, Bot.); Askerova in Fl. Azerb. VIII, 552; Kovalevskaja in Fl. Uzb. 

VI, 489. — Sonchus tataricus L. Mant. II (1771) 572. — Mulgedium 
runcinatum Cass, in Diet. sc. nat. XXXIII (1824) 296, p. p. — Agathyrsus 
tataricus D. Don in Edinb. New Phil. Joum. VI (1829) 310; Klok. in 
Vizn. Rosl. URSR, 603. — Sonchus maritimus auct non. L.: Glaus in 
Goebel, Reise II (1838) 284 (fide Ldb. 1. c, v. infra).— Mulgedium 
tataricum DC. Prodr. VII, I (1938) 248; Lbd. Fl. Ross. II, 2, 842; Boiss. 
Fl. or. Ill, 799; Fedtsch and Fler. Fl. Evrop. Ross. 1055; O. and B. 
Fedtsch. Perech. Rast. Turk. IV, 350 (Consp. Fl. Turk. No. 2857); Navski 
in Somye Rast SSSR IV, 339; Perf. Fl. Sev. Kraya II-III, 375; Kraschen. 
in Fl. Yugo-Vost. VI, 464; Govorukhin, Fl. Urala 521; Pavlov, Fl. Tsentr. 
Kazakhst. Ill, 361; Popov in Tr. Uzb. Gos. Univ. Nov. Ser. No. 27, Biol. 
Vyp. 14, 104; Kirp. in Majevski, Fl. (1954) 630; Popov, Fl. Sr. Sib. II, 
786; Nikitina, Mater, po Fl. Sev. Ski. Kirg. Ala-too 110, No. 1126.— M. 
tataricum a. typicum |3. floccosum and y. integrum Rgl. in Bull. Soc. Nat. 
Mosc. XI, II, 3 (1867) 181; O. and B. Fedtsch. op. cit. 361.— M. tataricum 
p. subfloccosum Rgl. in Tr. Peterb Bot. Sada I, 2 (1872) 255.— M 
tataricum var. anomalum Korsh. in Mem. Acad. Sc. Petersb. VIII ser. 

VII, 1 (1898) 264 (Tentam. Fl. Ross. or. No. 776).— M. tataricum f. 
pumilum B. Fedtsch. in Mat. Fl. Pam i Alaisk. Khr. (1900) 8 (Prilozh. 

283 from "Zemlevedeniyu", Kn. 1); O. and B. Fedtsch. op. cit. 351. — Cicerbita 
tatarica Beauverd ex Sosn. in Takht. and Fed. Fl. Erevana (1945) 338. — 
Lagedium tataricum Sojak in Novit. bot. Horti bot. Univ. Car. Prag. (1961) 
34.— Ic: Nevski. op. cit. 340, Fig. 492; Kraschen. op. cit. 465, Fig. 720; 
Hegi, op. cit, 1131, Fig. 807 — Exs.: Lang and Szovitz, Herb. ruth. No. 
144 (sub Soncho tatarico) Callier, Iter Taur. tert. (1900) No. 664 (sub 
Mulgedio tatarico). 



281 

Perennial. Plants with strong root system and vertical or slightly 
twisted rhizomes underground, producing aerial shoots. Stem (5)30- 
80(100) cm, according to published reports up to 190 cm high, sUghtly 
sulcate, glabrous or sometimes partly tomentose, or with isolated hairs, 
branched usually only above, or, less often, more or less throughout. 
Leaves approximate in lower part of stem, glaucous or light glaucous, 
rather thick, fleshy, but somewhat stiff, narrowed toward base; lower leaves 
amplexicaul, petiolate, pinnatisect or runcinate, with narrow, deflexed, 
usually acuminate lobes, sparsely spinulose along margin; middle leaves 
like lower leaves (but smaller) or undivided, lanceolate; upper leaves 
sessile or subsessile, undivided lanceolate, or narrow-lanceolate; very rarely 
all leaves undivided. Capitula with about 20(16-23) florets, cylindrical or 
truncate-turbinate, 11-14(20) mm long with fruits, in paniculate 
inflorescence, on unequal peduncles, covered with small, scaly leaves. 
Involucral bracts usually four-rowed, very finely pubescent on outer 
surface, often sUghtly violet; inner bracts with membranous, usually sinuate 
border, subobtuse. Corolla lilac-blue or blue. Achenes 5.5-6 mm long 
and 0.75-1 mm wide, oblong-linear or ellipsoid, slightly appressed, 
yellowish-brown to dark olive or almost black, with five to seven 
prominent longitudinal ribs, rather densely covered with very short stiff 
hairs (under high magnification!), gradually attenuate at apex into beak- 
like structure, usually lighter colored than and one-third to half as long 
as achene, terminating in funnel-shaped disk bearing pappus of thin, white 
or dirty yellowish, soft and lustrous, almost smooth and rather fragile 
hairs, one and one-half to two times as long as achene. Flowering May 
to July(September); fruiting June to September. (Plate XVI, Fig. 4.) 

On clayey and sandy soils (including saline lands), on steep river 
banks, banks of water bodies (rivers, lakes, irrigation channels, near wells), 
coastal sands; from zone of mixed forests to deserts, but particularly in 
the southern steppes and semideserts; mountains up to 3,400 m; as a 
weed in fields of different cultivated crops, fallow land, kitchen gardens, 
melon fields, orchards and in forest protection zones, as well as on 
284 irrigated lands, in oases and near springs; often in large clusters (patches) 
all over. — European Part: Dvina-Pechora (western part, less often) 
Ladoga-Ilmen (very rarely, one specimen seen from vicinity of Ostrov), 
Upper Volga (less often), Volga-Kama (mainly southern part of western 
half), Volga-Don (less often), Trans-Volga Region (commonly), 
Bessarabia, Black Sea Region, Crimea, Lower Don, Lower Volga 
(commonly); Caucasus: all regions; Western? Siberia: Upper Tobol, Irtysh, 
Altai; Eastern Siberia: Angara- S ay ans; Soviet Central Asia: all regions. 
General disbribution: central Europe, Atlantic Europe, Balkans-Asia 
Minor, Iran Region (Afghanistan), ?Indo-Himalayas (northern India, 
literature records), Dzhungaria-Kashgaria, Mongolia, China, ?Tibet 



282 

(literature records). Also reported from Scandinavia (cf. Lindeman, Svensk 
fanerogamflora, Andra uppl. (1926) 559), but this is very doubtful. 
Described by Linnaeus from "Tartaria" and Siberia. Type in London. 

Note. L. tatarica is not uniform throughout its urde range; however, 
the variations are not important enough to warrant a higher rank than 
form or modification. First of all, let us take the achene. It has been 
reported in the literature that almost beakless forms (f. pumilum B. 
Fedtsch.) are found in L. tatarica. But the achenes are always 
underdeveloped in stunted plants of L. tatarica growing under extreme 
or generally^ unfavorable conditions, and in this species there is a general 
tendency toward suppressed reproduction from seeds. On the basis of 
this character S. Korshinsky described the new var. anomala Korsh. for 
the extensive material in which it was impossible to find developled 
achenes. The usually cited difference in the achenes of L. tatarica and L. 
sibirica is extremely undependable; quite often, the beak in the former 
species is not any longer than in the latter. These two species are 
distinguished better on the basis of their vegetative characters and biology 
and also by the length of the corolla tube, which is about 4 mm in L. 
sibirica and usually 6 mm in L. tatarica. 

The vegetative organs of L. tatarica differ slightly in their pubescence, 
but may have larger differences in size, consistency, and shape of leaves. 
It is_quite apparent that the leaves of many Central Asian plants are 
strongly glaucous and extremely stiff. The plants collected from the 
Caspian lowlands, Balkhash Region, vicinity of Panfilov, and, especially, 
the Kyzyl-Orda District are characterized by very long (in individual 
cases up to 35 cm), linear-lanceolate or linear cauline leaves. L. tatarica 
from these places needs special attention; it would be particularly important 
to study the biology of the Mediterranean representatives in comparison 
with their ancestral types, which are distribubed more to the west and 
north. 

Economic Importance. Tatar lettuce is one of the pernicious and 
difficult-to-eradicate, sucker-producing weeds. It usually reproduces 
vegetatively through numerous adventitious buds on the roots and 
regenerating buds on the rhizomes. It has a strong root system with the 
tap root reaching a length of up to 1 m in annuals, and 4-5 m in adult 
285 perennials. Large, more or less horizontal roots arise from the main root 
(called propagation roots, as most of the buds are particularly concentrated 
on them). A single plant of L. tatarica can produce more than 50 aerial 
shoots in a single vegetative period. Control measures for this weed 
include, besides the use of herbicides (to which it is fairly resistant, 
owing to the thick cuticle on the leaves), a whole system of agronomic 
practices. For details about the biology of this plant, cf. Sornye 
Rasteniya SSSR [Weeds of the USSR] (Vol. IV, Acad. Nauk SSSR, 



283 

Moscow-Leningrad, 1935), and especially the article by A.M. Bilova, 
"Moloken tatarskii" [Tatar lettuce] (in Biology of Weeds. Handbook 
for Teachers, Uchpedgiz, Moscow, 1960). Moreover, L. tatarica is of 
major importance as a fodder plant, although it is poisonous in 
large quantity (for details, refer to the book by Larin et al., Kormovye 
Rast. Senokosov i Pastbishch SSSR [Fodder Plants of Grasslands and 
Pastures of the USSR], Vol. Ill, Sel'khozgiz, Moscow-Leningrad, 1956, 
678 p.) 

3. L. winkleri Kirp. nom. nov. — Mulgedium longifolium C. Winkl. 
in Tr. Peterb. Bot. Sada, XI, 10 (1890) 337 (serosum impressum; in 
volumino completo XI, 2 (1892) 337); O. and B. Fedtsch. Perech. Rast. 
Turk. IV, 351 (Consp. Fl. Turk. No. 2858); Zakirov, Fl. i Rastit. Bass. 
Zeravsh. II, 426. 

Perennial. Roots ? Stem 60-100 cm high, solitary, fistular, distinctly 
but shallowly sulcate, glabrous or subglabrous, sparsely leafy, weakly 
branched above (sometimes from middle). Leaves broadly or oblong-linear 
to lanceolate, gradually reduced toward apex, broadly but finely erose 
along margin, unevenly toothed-spinose; lower and middle cauline leaves 
basally narrowed into winged petiole, semiamplexicaul, usually with 
prominent thick midrib, usually entire or sometimes more or less runcinate; 
upper cauline leaves sessile or subsessile, semiamplexicaul; peduncular 
leaves reduced, scaly. Capitula of about 15 florets, almost cylindrical or 
slightly broadened above, up to 15 mm long with fruits, in lax panicles 
or racemose-paniculate inflorescence; peduncles usually shorter than 
capitula. Involucral bracts usually three-rowed, with very thin and dense 
appressed pubescence on dorsal surface (under a lens!), somewhat 
squarrose. Corolla?. Achenes about 4.5 mm long and 0.8 mm wide, 
oblong-ellipsoid, slightly flattened, yellowish or sand-colored, with 
prominent longitudinal ribs, densely covered with very short light-colored 
hairs (under high magnification!), slightly narrowed in upper part, without 
distinct beak, terminating in narrow disk bearing pappus; pappus of 
numerous, very thin, silky, snow-white hairs, almost smooth (poorly 
286 developed barbs detected only under high magnification), slightly curved 
at base, easily detached individually, up to 7-9 long. Flowering September. 
(Plate XVI, Fig. 1.) 

In upper mountain zone ("tau"); shady habitats; on clayey-sandy, 
moist soil and wet stony places. — Soviet Central Asia: Pamiro-Alai. 
Endemic. Described from the Alai Valley (Bolshoi Karamuk) on the basis 
of collections of Nevski. Type and isotypes in Leningrad. 

Note. Described entirely from only three (albeit authentic) specimens 
collected without the lower part. According to K. Winkler, who described 
this species, it is closely related to L. tatarica, from which it differs by 



284 

having a beakless achene (almost the same as achenes of Sonchus), leaves 
almost two times as long, and a much taller stem. 

After transfer from the genus Mulgedium to the genus Lactuca, it is 
not possible to retain the epithet "longifolium." as Lactuca longifolia 
Michx. (1803) and L. longifolia DC. (1838) already exist. 

Subgenus 2. Lactucopsis (Sch. Bip.) Babcock, Stebbins and Jenkins 
in Cytologia, Fujii Jubil, vol. (1937) 191 p. p. — Lactucopsis Sch. Bip. in 
Vis, and Pane, in Mem. Reale Inst. Veneto, XV (1870) 5 (PI. Serb. rar. 
III). — Lactuca sect. 4. Cicerbita Benth. in Benth. and Hook. f. Gen. pi. 

II (1873) 525, p. min. p. — Lactuca § 2. Scariolae *Biennes Boiss. Fl. or. 

III (1875) 804, p. p. — Cicerbita § 2. Lactucopsis Beauverd in Buil. Soc. 
Bot. Geneve, 2 Ser. II (1910) 116 (seorsum impressum. Contrib. a I'etude 
des Compos. Ill, p. 18). — Achenes flattened, almost black, oblong- 
ellipsoid, with two to five strongly raised longitudinal ribs on each side, 
covered with stiff upward-directed hairs (under a lens!). Particularly 
noticeable along sides of upper narrowed part as well as by transverse 
sinuate rows of very short outgrowths or hairs (under high magnification!); 
achene apex attenuate, beak-like, concolorous with remaining part; pappus 
of white, rather thin, usually very fragile hairs. Corolla yellow. 

Mostly biennial plants with thick or often furcate root, growing in 
forest-steppe and steppe zones as well as in mountains (up to middle 
mountain zone) and forests, on forest edges and in thickets of shrubs. 

The beak in typical members of Lactuca is a unique morphological 
structure, whereas the so-called beak in the members of the subgenus 
Lactucopsis is simply the greatly extended upper part of achene. 

Type of subgenus: Lactuca chaixii Vill. 

288 4. L. stricta Waldst. and Kit. Descr. et ic. pi. rar. Hungar. I (1802) 

47; DC. Prodr. VII, 1, 138; Boiss. Fl. or. Ill (1875) 808 (incl. (3. armena 
Boiss.) Grossh. Fl. Kavk. IV (1934) 256 and Opred. Rast. Kavk. 516; 
Klok. in Vizn. Rosl. URSR, 604; Szafer, Kulcz., B. Pawl. Rogl. Polsk. 
(1953) 774; Stank, and Tal. Opred. Vyssh. Rast. (1957) 444. —Cicerbita 
corymbosa Walk. Sched. crit. I (1822) 434. — Lactuca stricta Waldst. and 
Kit. p. runcinata and y. heterophylla Bogenh. Taschenb. Fl. Jena (1850) 
269. — L. quercina a. pinnatifida Bisch. Beitr. z. Fl. Deutschl. Cichorieen 
(1851) 205.— L. quercina L. a. typica Fiori Nuova Fl. analit. Ital. II (1925) 
825. — L. quercina L. ssp. stricta (Waldst. and Kit.) Hayek in Hegi, 111. 
Fl. VI, 2 (1928) 1128; Dostal, Kle^ (1954) 802.— L. quercina auct. fl. 
Ross. p. p. non L.: Ldb. Fl. Ross. II, 2 (1954) 806; Schmalh. Fl. II, 168; 
Fedtsch. and Fler. Fl. Evrop. Ross. 1059. — Ic: Waldst. and Kit. op. cit. 
t. 48 (color., bona); Dostal, op. cit. 799, fig. 2220. 



285 




287 Plate XVm. 

1 — Lactuca altaica Fisch. and Mey., habit of plant and achene; 2 — L. serriola Tomer, 

achene; 3 — L. sativa L., achene; 4 — L. auriculata DC, achene; 5 — L. undulata Ldb., achene; 

6 — L. rosularis Boiss., achene; 7 — L georgica Grossh., achene; 8 — L. saligna L., achene; 

9 — L. glauciifolia Boiss., achene; 10 — L. spinidens Nevski, achene. 



286 

Biennial. Root often furcate. Stem (50)80-150(200) cm high, weakly 
sulcate, smooth or almost smooth. Leaves glabrous or (especially beneath) 
slightly appressed-hairy; lower leaves early- withering, lyrately pinnatisect, 
with large apical ovate segment, narrow-winged, semiamplexicaul, with 
petiole expanded at base; middle leaves sessile, auriculate-sagittate at base, 
pinnatipartite or pinnatisect with oblong, ovate, or linear-lanceolate, or 
irregular-in-shape, more or less coarse-toothed lobes or segments; upper 
leaves strongly reduced, narrowly triangular or linear, entire, acuminate, 
reduced in inflorescence. Capitula of about 10 florets, cylindrical or almost 
cylindrical, (10)11-13(14) mm long with fruits, in somewhat lax 
corymbose-paniculate inflorescence; peduncles slender, usually longer than 
capitulum, with few reduced leaves resembling involucral bracts. Involucre 
usually three-rowed, bracts sUghtly appressed-hairy (under a lens!), longest 
bracts (inner) with membranous border, sometimes with more or less 
numerous red spots along dorsal surface. Corolla yellow. Achenes 
(excluding beak-like part) about 5 mm long and 1.25 mm wide, oblong- 
ellipsoid, flattened, dark violet or blackish, with (two) five prominent- 
longitudinal ribs on each side and transverse sinuate rows of light-colored 
membranous outgrowths and hairs and distinct (under magnification!) stiff, 
upward-directed hairs along sides of upper attenuate part of achene, near 
its transition to thin, black, beak-like constriction, latter 2-2.5 mm long, 
terminating in disk bearing pappus of fine, white, silky, almost smooth 
(under high magnification, slightly barbed), about 5 mm-long, fragile hairs 
falling individually. Flowering June to November. (Plate XVII, Fig. 3.). 
289 Deciduous forests and forest edges — European Part: Middle Dnieper, 

Bessarabia, Black Sea Region, Crimea; Caucasus: ?Ciscaucasia. General 
distribution: Scandinavia, central Europe, western Mediterranean, Balkans- 
Asia Minor. Described from Hungary. Type, apparently, in Budapest. 

Economic Importance. The milky latex from the stems of L. stricta, 
the so-called lactucarium, which, hardens when exposed to the air, contains 
the somniferous substance lactucerin, which has an effect similar to opium. 

5. L. chaixii Vill. Prosp. hist. pi. Dauph. (1779) 33 (non vidi!); Ej. 
Hist. pi. Dauph. Ill (1789) 154; DC. Prodr. VII, 1, 138 (ut species dubia); 
Grossh. Fl. Kavk. IV (1934) 256 and Opred. Rast. Kavk. 516; Klok. in 
vizn. Rosl. URSR 604; Schchian in Fl. Gruzii VIII, 684, Stank, and Tal. 
Opred Vyssh. Rast. (1957) 443. — Lactuca sagittate Waldst. and Kit. Descr. 
et ic. pi. rar. Hungar. I (1802) 1; DC. Prodr. VII, 1, 136; Ldb. R. Ross. 
II, 2, 804; Boiss. Fl. or: III, 807; Kraschen, in Fl. Yugo-Vost. VI, 471.— 
Prenanthes altissima Pall. Reise III (1776) 655 nomen. — Lactuca altissima 
M.B. Fl. taur.-cauc. II (1808) 242 and III (1819) 528; DC. Prodr. VII, 1, 
136; Ldb. H. Ross. II, 2, 805; Kirp. in Majevskii, Fl. (1954) 563.— L. 
stricta 6. integrifolia Bogenh. Taschenb. Fl. Jena (1850) 269. — L quercina 



287 

p. integrifolia Bisch Beitr. z. Fl. Deutschl., Cichorieen (1851) 206.— 
Lactucopsis chaixii and L. altissima Sch. Bip. in Vis. and Pane in Mem. 
Reale Inst. Veneto XV (1870) 6 (PI. Serb. rar. III). — L. quercina ssp. 
sagittata (Waldst. and Kit.) Soo. sec. Javorka. — Soo. A. Magyar Novenyv. 
Kez. II (1951) 723 Dostal, Klec (1954) 802.— L. quercina ssp. chaixii 
(Vill.) Hayek in Hegi, 111 Fl. VI, 2 (1928). 1128.— L. quercina auct. Fl. 
Ross. p. p. non L..: Lbd. Fl. Ross. II, 2, 806; Schmalh. H. II, 168; Fedtsch. 
and Fler. Fl. Evrop. Ross. 1059.— Ic: Vill. op. cit. (1789), t. XXXII; 
Waldst. and Kit. op. cit. t. 1 (color, bona, sub L. sagittata); Bonnier, Fl. 
Compl. 111. France, Suisse et Belg. VI (1923), t. 351, fig. 1682 (color.); 
Javorka and Csapody, Iconogr. Fl. Hungar. (1933), t. 561, fig. 4027. — 
Exs.: F. Schultz Herb. norm. nov. ser. cent. 19, No, 1811. 

Biennial. Root thick, narrow-conical, often furcate. Stem 60-100(150) 
cm high, weakly sulcate, fistular, smooth or almost smooth. Lower leaves 
obovate, narrowed toward base into petiole, early-withering; middle cauline 
leaves oblong-elliptical to lanceolate, sessile, semiamplexicaul, sagittate 
at base, spinulose-toothed or subentire; upper leaves reduced, lanceolate- 
linear. Capitula with about 10 florets, cylindrical or almost cylindrical, 
with fruits 10-13 mm long, in corymbs or corymbose-paniculate 
inflorescence; peduncles slender, usually longer than capitula, with few 
reduced leaves resembling outer involucral bracts. Involucre three- or four- 
rowed; involucral bracts lanceolate or linear; young bracts finely appressed- 
hairy on outer surface, mature bracts slightly pubescent (under a lens!); 
inner bracts longest, with membranous border, subobtuse, usually with 
290 numerous red spots on dorsal side. Corolla yellow. Achenes (excluding 
beak-like part) 4-5 mm long and 1.25 mm wide, oblong-ellipsoid, flattened, 
almost black, with (two) three to five prominent longitudinal ribs on each 
side, and transversely sinuate rows of light-colored membranous 
outgrowths or somewhat stiff hairs (under a hand lens!), apically narrowed 
into black, very thin beak-like part, up to 2.5 mm long and terminating in 
light-colored disk bearing pappus; pappus of very fine, white, silky hairs, 
almost smooth (under high magnification slightly barbed), fragile, falling 
individually, 5-6 mm long. Flowering June to August. 

Deciduous forests and shrub thickets; forest-steppe and steppe zones, 
in the Caucasus up to middle mountain zone, — European Part: Upper 
Dnieper, Middle Dnieper, Volga-Don, Trans- Volga Region, Black Sea 
Region, Crimea, Lower Don; Caucasus: Ciscaucasia, Eastern 
Transcaucasia, ?Westem Transcaucasia (Novorossisk, literature records, 
?southem Transcaucasia (rarely; literature records). General distribution: 
central Europe, Atlantic Europe, western Mediterranean, Balkans-Asia 
Minor. Described from southeastern part of France. Type in Paris(?). 

Economic Importance. The milky latex has properties similar to 
that of L. stricta (cf. p. 286) 



288 

6. L. wilhelmsiana Fisch. and Mey. in DC. Prodr. VII, 1 (1838) 
134; Ldb. Fl. Ross. II, 2, 803; Boiss. Fl. or. Ill, 808; Grossh. Fl. Kavk. 
IV (1934) 257 and Opred. Rask. Kavk. 516; Klok. in Vizn. Rosl. URSR 
604; Schchian in Fl. Gruzii VIII, 687; Stank, and Tal. Opred. Vyssh. 
Rast. (1957) 443; Askerova in Fl. Azerb. VIII, 553.— L. stricta Waldst. 
and Kit. var. Hohenack. in Bull. Soc. Nat. Mosc. XI, 3 (1838) 270.— 
Lactucopsis wilhelmsiana Sch. Bip. in Vis. and Pane, in Mem. Reale Inst. 
Veneto XV (1870) 6 (PL Serb. rar. III).— Exs.: PI. or. exs. No. 75. 

Biennial. Root often tuberous. Stems (50)60-120(150) cm high, 
weakly sulcate, fistular, smooth or almost smooth. Leaves (excluding 
upper) pinnatisect, semiamplexicaul, with short auriculate-sagittate base, 
lower leaves petiolate, other sessile; leaf segments narrow-triangular or 
linear-lanceolate, less often irregular in shape, often antrorse or retrorse, 
sometimes erect and horizonteil, entire or more or less distinctly spinose- 
toothed; upper cauline leaves narrow-triangular or linear, reduced in 
inflorescence. Capitula of about 15 florets, cyUndrical or almost cyUndrical, 
with fruits (10)13-15 mm long, in lax, often few-headed corymbose- 
paniculate inflorescence; peduncles slender, unequal, with one to few 
reduced leaves, resembling outer involucral bracts. Involucre three- or 
four-rowed; young involucral bracts finely appressed-hairy on outer 
291 surface with very short papilliform hairs, mature bracts weakly pubescent 
(under a hand lens!); inner bracts longest, with membranous border, 
apically attenuate, often with more or less numerous red spots on dorsal 
surface. Corolla yellow. Achenes (excluding beak-like part) about 5 mm 
long and 1.25 nun wide, oblong-ellipsoid, flattened, or almost so, with 
five or more prominent longitudinal ribs on each side, and transversely 
sinuate rows of light-colored membranous outgrowths or stiff hairs (under 
a lens!) most distinct in upper narrowed part of achene, very gradually 
extended into thin, black, beak-like part, up to about 3.5 mm long, 
terminating in light-colored disk bearing pappus; pappus of very fine white, 
silky, almost smooth hairs (under high magnification, slightly barbed), 
fragile, falling individually, 5-6 mm long. Flowering May to August. 
(Plate XVII, Fig. 1.) 

Steppe, forest-steppe, and middle mountain zone; edges of deciduous 
forests, in shrub thickets. — European Part: Middle Dnieper. Black Sea 
Region (both regions of the European USSR, indicated only on the basis 
of literature records and needing confirmation); Caucasus: Ciscaucasia, 
Eastern and Southern Transcaucasia, Endemic (?). Described from territory 
of present Nakhichevan ASSR. Type in Leningrad. 

Note. This species is rather doubtful, and its separate status needs to 
be confirmed. It is often mentioned in descriptions that the "beak" (i.e. 
the narrow beak-like part) in L. wilhelmsiana is as long as the achene. 
This is not correct, as the narrowed part itself almost never exceeds 3 



289 

mm in length. In practice, the fruits of L. wilhelmsiana can be distinguished 
from those of L striata only with difficulty. The tuberous thickening of 
the root, also given as a characteristic feature of L. wilhelmsiana is not 
always developed. Doubt about the separateness of Lwilhelmsiana was 
raised already by Boissier (I.e.) in his note on this species. 

Subgenus 3. Pterachaenium (Kitam.) Kirp. stat. nov. — Lactuca sect. 
Pterachaenium Kitam. in Acta Phytotax et Geobot. VI (1937) 237. 

Achene completely flat, almost black, ellipsoid or oblong-ellipsoid, 
with one or few prominent longitudinal ribs, appearing slightly rugose 
from transversely sinuate rows of very short hairs (under high 
magnification!), apically attenuate into small conical neck with very short 
yellowish-green beak, or terminating in short but broad collar, without 
beak, and abruptly transitional to broadly truncate, funnel-shaped disk 
bearing pappus. Pappus of white, fine, fragile hairs, falling individually. 
Corolla yellow or pale yellow. 

Biennial or perennial, mostly forest plants, with characteristic 
292 underground, narrow, fusiform thickenings. Only a few species are 
indigenous to the USSR, growing almost exclusively in the Far East and, 
less frequently, in Eastern Siberia; outside the USSR, in the Korean 
Peninsula, China, Japan, India, Indochina, Sunda Islands and the 
Philippines. The subgenus is botanically and geographically quite isolated 
from the other subgenera of Lactuca, and less closely related to typical 
Lactuca. Type of subgenus : Lactuca indica L. 

7. L. indica L. Mant. II (1771) 278; DC. Prodr. VII, 1,136; Gagnep. 
in Fl. Gen. Indo-Chine III, 654; Ling in Contrib. Inst. Bot. Nat. Acad. 
Peip. Ill, 4, 187; Kitag. in Report Inst. Sc. Res. Manch. Ill, App. 1, 456 
(Lineam. fl. Manshur.); Kitam. in Mem. Coll. Sc. Univ. Kyoto, ser. B, 
XXIII, 1 (1956) 137, incl. f. runcinata Kitam. (Compos. Jap. V). — 
Prenanthes squarrosa Thunb. Fl. Jap. (1784) 303. — Lactuca brevirostris 
Champ, ex. Benth. in Hooker's Joum. Bot. IV (1852) 237; Hook, f Fl. 
Brit. Ind. Ill, 405. — L. amurensis Rgl. and Maxim, ex Rgl. in Ind. Sem. 
Hort. Petrop. (1857) 42; Maxim, in Mem. Acad. Petersb. div. sav. IX, 
178 (Primit. Fl. Amur. No. 457). — L. squarrosa Miq. in Ann. Mus. Bot. 
Lugd.-Batav. II (1866)189; Ej. Prolus. Fl. Jap. (1866-1867) 121 and 362; 
Maxim, in Bull. Acad. Sc. Petersb. XIX (1874) 526 (incl. f. runcinata 
Maxim, and f. indivisa Maxim.); Kom. in Tr. Peterb. Bot. Sada XXV 
(1907) 778 (Fl. Man'chzh. Ill, No. 1648) (incl. var. integrifolia Kom., 
var. dentata Kom. and var. runcinato-pinnatifida Kom. op. cit p. 779); 
Kom. and Alls. Opred. Rast. Dal'nevost. Kr. II, 1098, 1101; Nevski in 
Som. Rast. SSSR. IV, 344; Popov, Fl. Sr. Sib. II, 787.— L. indica f 
indivisa (Maxim.) Hara, Enum. Spermat. Jap. II (1952) 220. 



290 

Biennial. Main root usually robust, or plant with distinct, underground, 
narrow, fusiform, tuberous thickenings. Stem (60)90-120(200) cm high, 
often reddish (especially below), weakly sulcate, smooth, branched above. 
Leaves usually sessile, with long cusp or acuminate; all leaves, except 
uppermost, with somewhat thick midrib, often with narrow squamiform 
hairs, sometimes rather long, reddish; shape of leaves extremely variable: 
from entire, linear, oblong-linear, or linear-lanceolate, with 
semiamplexicaul auriculate-sagittate base (f. indivisa) to more or less strong 
pinnatisect or runcinate-pinnatisect, with sinuate or spinose-toothed 
segments (f. runcinata); lower leaves withering before anthesis, petiolate; 
uppermost leaves strongly reduced, linear-lanceolate or linear. Capitula 
of about 25 florets, almost cylindrical or broadly cylindrical, 13-15 nrmi 
long with fruits, usually in narrow paniculate or racemose-paniculate 
inflorescence; peduncles slender, unequal, but often longer than capitula, 
with few small, reduced, leaves resembling involucral bracts. Involucre 
293 mostly three-rowed; involucral bracts often green, very finely appressed- 
hairy on dorsal surface (under a lens!), more or less reddish at apex; at 
least inner bracts with membranous border. Corolla, when fresh (according 
to literature records), pale yellow or almost white, pale blue on drying. 
Achenes ellipsoid, about 5 mm long and 2-2.5 mm wide, sometimes 
slightly asymmetrical, almost black, flat, usually with only one prominent 
rib on each side and numerous light-colored stiff, upward-directed, 
very short (under a lens!) hairs, in sinuate-transverse rows throughout, 
apically attenuate into conical neck with short (about 1 mm long) 
yellowish-green beak, terminating in broad disk with more or less well- 
developed ridge bearing pappus; pappus hairs very thin, fragile, about 
8 mm long. Flowering (June) July to August (September). (Plate XVII, 
Fig. 4.) 

Old fields, weed in crops, riverine meadows and shrub thickets, 
meadow patches in open forests, rarely along sea coasts. — Eastern Siberia: 
Dauria (less often); Far East : Zeya-Bureya, Ussuri. General distribution: 
Japan, China, Indo-Himalayas. Besides, found in the Philippines, Java 
and Sumatra. Described from Java from collections of Osbeck. Type in 
London. 

Note. Apparently, the semi-weedy nature of this species explains its 
great variability. Particularly noticeable is the variability in leaf form. 
The most widespread variants are f. indivisa, with entire leaves, and f. 
runcinata, with more or less strongly divided, often runcinate leaves. The 
numerous other transitional forms do not deserve special names. 
According to V.L. Komarov, forms with entire leaves are confined to the 
meadows and shrub thickets with the drier soils, while the forms with 
divided leaves are found on the fertile soils. 



291 

So far, I have not been able to examine the authentic specimens. 
Nevertheless, the viewpoint of Kitamura, Merrill, Hanpen, Ling-Zhung 
and many others, identifying Prenanthes squarrosa Thunb. with Lactuca 
indica L., apparently is correct. The relatively southern collection of L. 
indica, which was available to us (for example, from southern China 
right up to Hainan Island), is characterized by entire leaves, sometimes 
flocculose-pubescent stems (under a lens!) and a somewhat strongly 
developed ridge on the pappus-bearing disk. However, considering the 
extreme polymorphism of this species, these differences can hardly be 
accorded taxonomic significance. 

In the Herbarium of the Botanical Institute, Academy of Sciences of 
the USSR, the two sheets collected by Stukov from die Transbaikal Region 
were preserved under the name ""Mulgedium floribundum Zing." I did 
not find any description under this name. Hence, these specimens should 
be included under L. indica. 

8. L. triangulata Maxim, in Mem. Acad. Petersb. div. sav. IX (1859) 
177 (Primit. Fl. Amur. No. 457); Rgl. in Mem. Acad. Petersb. ser. VII, 
4, 97 (Tentam. Fl. Ussur. No. 306); Kom. in Tr. Peterb. Bot. Sada, XXV, 
294 777 (Fl. Man'chzh. Ill, No. 1647); Kom. and Alls. Opred. Rast. 
Dal'nevost. Kr. II, 1098; Ling in Contrib. Inst. Bot. Nat. Acad. Peip. Ill, 
4, 187; Kitag. in Report Inst. Sc. Res. Manch. Ill, App. 1, 456 (Lineam. 
fl. Manchur.); Sugaw. 111. fl. Saghal. IV, 1931; Kitam. in Mem. Coll. Sc. 
Univ. Kyoto, ser. B, XXIII, 1, 142 (Compos. Jap. V); Vorobev in Tr. 
Dal'nevost. Fil. Ser. Bot. III(V), 77 (Materialy k Fl. Kuril' skikh o-vov. 
No. 975). — L. triangulata Maxim, var. sachalinensis Kitam. in Acta 
Phytotax. et Geobot. XI, 2 (1942) 126.— Ic.:Sugaw. op. cit. t. 889. 

Biennial or pemnial. Plants with more or less distinct, underground, 
narrow-fusiform, tuberous thickenings. Stem 60-120(250) cm high, 
shallow-sulcate, green or reddish below, branched from middle or, often, 
from upper third (and above). Leaves thin, green above, glaucous beneath, 
unevenly spinose-toothed along margin, sometimes with few, narrow, pale, 
squamiform hairs (under a lens!) also on surface and between teeth; lower 
leaves (usually early-withering and therefore frequently absent) on long, 
narrow-winged, slightly expanded at base, almost semi-amplexicaul 
petioles, terminating in almost half as long, triangular, triangular-ovate, 
or triangular-hastate lamina, acute, emarginate-reniform at base; petioles 
of middle cauline leaves gradually reduced, but increasingly broad-winged, 
with broad, hastate or auriculate, semiamplexicaul base and usually 
triangular or rhombic lamina; upper cauline leaves reduced, sessile, 
eUiptical to oblong and lanceolate. Capitula, with 10-15 florets, cylindrical 
or cylindrical-campanulate, about 10 mm long with fruits, in rather lax 



292 

and narrow-paniculate or racemose-paniculate inflorescence; peduncles 
slender, conical, usually much longer than capitula, slightly swollen above 
below capitulum. Involucre two- or three-rowed; involucral bracts with 
fine appressed pubescence on dorsal surface (under a hand lens!); inner 
bracts longer, slightly narrowed in middle, subobtuse, narrowly 
membranous along margin. Corolla yellow. Achenes about 4 mm long 
and 2 mm wide, ellipsoid or broadly oval, flat or almost flat, sometimes 
concave, dark brick-red to dark violet and almost black, with one strongly 
raised longitudinal rib each side, or often two on one side; achenes (under 
a hand lens!) appear rugose or divided into individual cells from transverse 
sinuate rows of very short, upward-directed hairs (more distinct on lateral 
sides of achene), apically shallow-emarginate, bearing short (0.2-0.3 nmi 
long) and relatively broad collar from center terminating into still broader 
light-colored annulus bearing about four- or five-angled disk with pappus 
of very thin, fragile, 6-7 mm-long hairs. Flowering July to August. (Plate 
XVII, Fig. 5.) 
295 In shady mountain forests, on stony but humus-rich soils; less often 

in herb meadows, on gravel deposits of mountain streams, etc. — Far East: 
Ussuri, Sakhalin (Sakhalin, Iturup Island among the Kurils). General 
distribution: Japan (Hondo Island). Korean Peninsula, northeastern China. 
Described from collections of Maximowicz from Amur. Lectotype and 
paratypes preserved in Leningrad. 

Note. Kitamura recognized L. triangulata Maxim, var. sachalinensis 
Kitam., with a crisped-hairy stem. This character was not found in the 
scanty material at our disposal. 

9. L. raddeana Maxim, in Bull. Acad. Sc. Petersb. XIX (1874) 526 
and in Mel. Biol. IX, 3 (1874) 355; Kom. in Tr. Peterb. Bot. Sada XXV, 
776 (Fl. Man'chzh. Ill, No. 1646); Kom. and Alls. Opred. Rast. 
Dal'nevost. Kr. II, 1098; Ling in Contrib. Bot. Nat. Acad. Peip. Ill, 4, 
188; Kitag. in Report Inst. Sc. Res. Manch. Ill, App. 1, 456 (Lineam. fl. 
Manshur.); Sugaw. 111. fl Saghal. IV, 1929; Kitam. in Mem. Coll. Sc. 
Univ. Kyoto, ser. B, XXIII, 1, 138 (Compos. Jap. V); Vorobiev in Tr. 
Dal'nevost. Fil. Ser. Bot. III(V), 77 (Materialy k Fl. Kuril' skikh o-vov, 
No. 974). — L amurensis auct. non Rgl. and Maxim.: Herd, in Bull. Soc. 
Nat. Mosc. XLIII, I, 2, 109 (PI. Raddeanae Monopet. IV, No. 222).— L. 
triangulata auct. non Maxim.: Herd. ibid. (No. 223). — Ic: Sugaw. op. 
cit. t. 888. 

Biennial (or perennial?). Plants with more or less distinct, 
underground, narrow-fusiform thickenings. Stem 65-120(200) cm high, 
weakly sulcate, often reddish, usually rather densely covered with narrow 
squamiform hairs, branched above and glabrous here. Leaves highly 
variable, green above, glaucous beneath, unevenly erose, spinose-toothed, 



293 

and usually with narrow reddish frill, and short squamiform hairs (under 
a hand lens!) on both sides and along margin (more distinct along veins 
beneath); lower leaves (early- withering) mostly on long, narrow- winged 
petioles, with lyrate-pinnatisect (or pinnatipartite) lamina, with triangular- 
hastate or triangular-ovate apical segment, and one to three pairs of lateral 
segments usually retrorse; middle and upper cauline leaves on broad- 
winged petioles, sometimes with a pair of lateral lobes, often without, 
with ovate, ovate-triangular or oblong-rhombic lamina; uppermost leaves 
strongly reduced, sessile, mostly oblong or lanceolate. Capitula with 9- 
10 (according to literature records, up to 14-15) florets, almost cyUndrical, 
slightly swollen in lower part and somewhat broadened in upper, about 
10 mm long with fruits, usually in narrow-paniculate inflorescence; 
peduncles slender, unequal, but often as long as capitula or longer, with 
few reduced leaves resembling involucral bracts. Involucre 
296 mostly two-rowed; involucral bracts herbaceous; inner bracts longer, 
narrow-ovate, often reddish, more or less distinctly narrowed below 
middle, convex, subobtuse. Corolla (according to literature records) pale 
yellow or straw-yellow. Achenes 3-4 mm long, 1.5-2 mm wide, ellipsoid, 
flat, often curved, dark violet to almost black, with few strongly raised 
ribs on each side and light-colored short hairs in transverse sinuate-rows, 
more distinct in upper part of achene (under a lens!), toward base slightly 
narrowed, with very short (sometimes not distinct) 0.1-0.3 mm long collar 
at apex, terminating in light colored, roundish, four- or five-angled disk 
with pappus; pappus of thin, about 6 mm-long, fragile hairs, falling singly. 
Flowering June to September. (Plate XVII, Fig. 2.) 

In valley meadows among tall grass, shrub thickets, logged areas, 
deciduous (mainly open) forests, very rarely in old fields. — Far East: 
Zeya-Bureya, Ussuri, Sakhalin (Sakhalin and southern Kurils). General 
distribution: Japan, Korean Peninsula, northeastern China. Described by 
Maximowicz from Amur Basin. Lectotype (collections of Radde from 
Bureya Range) and paratypes preserved in Leningrad. 

Note. Kitamura (Joum. Jap. Bot. XXI (1947) 52) recognized L. 
raddeana var. elata (Hemsl.) Kitam. (basionym: L. elata Hemsl. in Joum. 
Linn. Soc. XXIII (1888) 481). It differs from the typical form by having 
a tall (up to 2 m) habit and usually three-ribbed achene. Kitamura reported 
it from Sakhalin and Kunashir. There are only two sheets from Sakhalin 
in the Herbarium of the Botanical Institute, Academy of Sciences of the 
USSR, while there are no collections at all from Kunashir. This variety is 
not apparent on the basis of the material examined by us from the USSR. 
The question of var. elata remains unresolved for the present. 

L. raddeana is a species with extremely variable leaves; the major 
forms are mentioned in the description. Besides, plants are found in which 
the leaves are sessile from the very base or almost from the base of the 



294 

stem and have an entire or nearly entire lamina; the upper and middle 
leaves are sometimes pinnatilobate or pinnatipartite, or they have a 
rhomboid, ovate, or oblong lamina (sometimes also a different form of 
the lamina) with a more or less broad, cuneately winged petiole. These 
variations in leaf shape have no taxonomic importance. 

The ranges of L. raddeana and L. triangulata partly coincide. However, 
intermediate or hybrid forms have not been found in the contact zone 
(probably because of the significant ecological isolation of these species). 

Subgenus 4. Lactuca. — Lactuca sect. I. Scariola DC. Prodr. VII, 1 
(1838) 133, p. p. — Lactuca subgen. Scariola Babcock, Stebbins. and 
Jenkins in Cytologia, Fujii Jubil. vol. (1937) 191, p. p. solum nomen. 

297 Achenes flattened or flat, variable in color (gray, brownish, dark 

brown of different shades, olive, dark violet to almost black), narrow- 
obovoid or oblong-ellipsoid, with rather numerous or, less often, only 
one to three prominent longitudinal ribs, usually covered with short, erect, 
light-colored hairs (under a hand lens!); besides, achene surface often 
with very short, transversely sinuate stripes of extremely short hairs (under 
high magnification!); a thin filiform beak at achene apex, generally 
differing in color from rest of achene. Pappus of white, very thin hairs, 
fragile or more or less strongly attached to the broad disk. Corolla yellow, 
less often lilac or pink. 

Annual or biennial, less often perennial herbs or semishrubs, mostly 
growing in dry (steppe, semidesert, or desert) regions of somewhat saline 
soils. 

Type of subgenus: type of genus. 

Section 1. Lactuca. — Lactuca sect. 1. Scariola § 2. Xanthinae DC. 
Prodr. VII, 1 (1838) 135, p. p. — Lactuca sect. 1. Lactucae genuinae W.D. 
Kock, Synops. fl. Germ, et Helvet. (1837) 431, p. p. 

Achenes narrow-obovoid or oblong-ellipsoid, with more or less 
numerous prominent longitudinal ribs and well-developed filiform beak 
almost as long or one and one-half to two times as long as achene. Corolla 
yellow. 

Type of section: type of subgenus. 

10. L. serriola Tomer in L. Centur. II Plant. (1756) 29 (non vidi!) 
and in Amoenit. acad. IV (1759) 328; Grossh. Fl. Kavk. IV (1934) 257 
and Opred. Rast. Kavk. 517; Kraschen. in Fl. Yugo-Vost. VI, 469; Klok. 
in Vizn. Rosl. URSR (1950) 603; Schchian in H. Gruzil VIII, 688; Szafer, 
Kulcz., B. Pawl. Rosl. Polskie (1953)743; Kirp. in Majevski. Fl. (1954) 
633; Hayek in Hegi, 111. Fl. 2 Aufl. VI, 2, 1116; Dostal, Kli^ (1954) 801; 
Stank, and Tal. Opred. Vyssh. Rast. (1957) 443; Mikhailovskaja in Fl. 
SSSR V, 191; Kirp. in Fl. Turkm. VII, 335; Askerova in Fl. Azerb. VIII, 



295 

555; Kovalevskaja in Fl. Uzb. VI, 484. — L. scariola L. Sp. pi. ed. 2. 
(1763) 1119; DC. Prodr. VIII, 1, 137; Ldb. Fl. Ross. II, 2, 805; Schmalh. 
Fl. II, 168; Fedtsch. and Fler. Fl. Evrop. Ross. 1058; O. and B. Fedtsch. 
Perech. Rast. Turk. IV, 320 (Consp. Fl. Turk. No. 2783): Nevski in Som. 
Rast. SSSR IV, 341; Ling in Contrib. Inst. Bot. Nat. Acad. Peip. Ill, 4, 
190; Perf. Fl. Sev. Kraya II-III, 376; Govorukhin. Fl. Urala 520; Pavlov, 
Fl. Tsentr. Kazakhst. Ill, 366; Popov in Tr. Uzb. Gos. Univ. Nov. Ser. 
No. 27, Biol. Vyp. 14, 103; Kryl. Fl. Zap. Sib. XI, 3026; Grubov, Consp. 
Fl. MNR 282, No. 1856 (Tr. Mong. Kom., Vyp. 67); Kitam. in Res. 
Kyoto Univ. Exp. Karak. a. Hinduk. 1955, II, 431 (Fl. of Afghanistan, 
1960); Nikitina, Mater, po fl. Sev. Ski. Kirg. Ala-too 111, No. 1128; 
298 Vvedensky et al. Opred. Rast. Golodnoi Stepi 40 (Tr. Tashk. Gos. Univ. 
Vysh. 178, Bot.).— L. sylvestris Lam. Fl. Fr. II (1778) 84.— L. latifolia 
Gilib. Fl. Lithuan. Ill (1782) 234; Ej. Exerc. phytol. I (1792) 188.— L. 
virosa Luce, Topogr. Nachr. Ins. Oesel (1823) 260, non L. — L scariola. 
p. integrifolia Bogenh. Taschenb. Fl. Jena (1850) 269. — L scariola a. 
sylvestris Bisch. a. vulgaris and (3. integrifolia Bisch. Beitr. z. Fl. Deutschl. 
Cichorieen (1851) 189.— L. dubia Jord. Pugill. plant, nov. (1852) 119 
(non vidi!). — L. scariola j3. angustana Lindem. in Bull. Soc. Nat. Mosc. 
XLV, I, 2 (1872) 303.— Ic: Rchb.f. Ic. fl. Germ. et. Halvet. XIX (1859) 
t. 70 (MCCCCXXI) fig. 1-6 (sub L scariola); Javorka and Caspody, 
Iconogr. fl. Hungar. (1933) t. 561, fig. 4030; Hegi, op. cit. 1116, fig. 
795; Dostal, op. cit. 799; fig. 2216; Mikhailovskaja, op. cit. 193, Plate 
LXV.— Exs.: Kar. and Kir. No. 874; Callier, It, Taur. tert. (1900) No. 
792 (sub Lactuca scariola L. var. integrifolia Bisch.); Novopokr. Gerb. 
Donsk.Fl. No. 80; Hayek. Fl. Stir. exs. No. 1277; Billot, Fl. Gall. et. 
Germ. exs. No. 2704 (sub L. scariola L., sicut omnia exsiccata sequentia); 
Kickxia Belgica, No. 381; Petrak Fl, Bohem. et Morav. exs. No. 400; Fl. 
exs. austro-hung. No. 3398; Fl. exs. reip. Bohem. Slov. Nos. 634, 983. 
Annual or biennial. Stem (30)60-125 cm high, more or less sulcate, 
whitish or yellowish (sometimes weakly reddish-violet at base), smooth 
or with sparse stiff spinules below, branched in upper half. Leaves 
glaucous, erose-pinnatifid, often with two retrorse lobes, sessile, 
amplexicaul, with sagittate base, usually with yellowish stiff spinules along 
midrib beneath, spinulose-toothed, vertical in clear weather at sunny places; 
uppermost cauline leaves usually lanceolate, undivided; sometimes all 
leaves more or less undivided, densely spinulose-toothed. Capitula 
cylindrical or oblong, 10-13 mm long with fruits, with 15-25 florets, on 
peduncles in paniculate, often pyramidal inflorescence; peduncles usually 
longer than capitula, usually with one or two very small scaly leaves, 
often tomentose or papulose on outer surface, sometimes indistinctly 
transitional to involucral bracts. Involucral bracts deflexed on maturation 
of achenes, three- or four-rowed, arachnoid-lanate or papulose on outer 



296 

surface, usually with one to several violet spots or slightly violet, 
subobtuse, and sometimes with apical tuft of transparent hairs; outer bracts 
ovate or triangular; inner ones linear- or oblong-lanceolate, with 
membranous, slightly sinuate border. Corolla yellow (turning blue on 
drying). Achenes 3-3.5 mm long and up to 1 mm wide, gray, brownish, or 
olive-colored, narrowly obovoid or oblong-ellipsoid, flattened, with (five) 
seven to nine prominent longitudinal ribs on each side, along them with 
rather long, erect, light-colored hairs (under a hand lens!), mostly in upper 
299 part, and indistinct transversely sinuate stripes of fine hairs (under high 
magnification!) in between them; achene apices attenuate into about 0.5 
mm long, thin collar, converging in filiform whitish beak slightly or one 
and one-half (two) times as long as achene, terminating in flat-conical, 
reddish-brown disk bearing pappus of very thin, white, soft, slightly barbed 
(under a hand lens!), about 6 mm-long hairs. Flowering June to September 
(October). (Plate XVIII, Fig. 2.) 

Weedy places, near dwellings and roadsides, in crops and old fields, 
orchards, kitchen gardens and vineyards, on irrigated lands and near 
irrigation channels as well as in gorges, ravines and valleys in mountains 
and foothills near streams. — European Part: Dvina-Pechora (extreme 
southwest!), Baltic Region, Upper Volga, Volga-Kama, Upper Dnieper, 
Middle Dnieper, Volga-Don, Trans-Volga Region, Upper Dniester, 
Bessarabia, Black Sea Region, Crimea, Lower Don, Lower Volga; 
Caucasus: all regions. Western Siberia: Upper Tobol, Irtysh, Altai; Eastern 
Siberia: Angara-Sayans (south); Soviet Central Asia: all regions. General 
distribution: Scandinavia, central Europe, Atlantic Europe, Mediterranean, 
Balkans-Asia Minor, Armenia and Kurdistan, Iran, Indo-Himalayas, 
Dzhungaria-Kashgaria, Mongolia, China (northern), ?Beringia, ?Tibet, 
introduced in North America. Described from southern Europe. Type in 
London. 

Note. A highly variable species with numerous poorly delineated 
separate forms within its wide distribution range. In the European USSR, 
forms with more or less entire leaves are often found in Ciscaucasia and 
the northern Caucasus. Forms entirely without spinules are often found 
in Central Asia. Krascheninnikov labeled one such sheet as "Lactuca 
scariola L. v. laevissima Krasch." Quite sclerophyllous forms are found 
in the Nakhichevan ASSR as well as in the south of Central Asia, which 
differ by having stiff, flat and more glaucous leaves, and certain other 
features; however, they are too insignificant to be given serious taxonomic 
importance. 

Economic Importance. L. serriola has high fodder value. For details 
see Larin et al. Kormovye rastenlya senokosov i pastbishch [Fodder Plants 
of Grasslands and Pastures of the USSR], Vol. Ill, Sel'khozgiz, Moscow- 
Leningrad, 1956, 676 p. 



297 

ILL. sativa L. Sp. pi. (1753) 795 (incl. p. capitata L. and y. crispa 
L.); DC. Prodr. VII, 1, 138; Ldb. Fl. Ross. II, 2, 806; Schmalh. Fl. II, 
168; Kom. in Tr. Peterb. Bot. Sada XXV, 780 (Fl. Man'chzh. Ill, sine 
numero); Grossh. Fl. Kavk. IV (1934) 258 and Opred. Rast. Kavk. 517; 
Kraschen. in Fl. Yugo-Vost. VI, 469; Perf. Fl. Sev. Kraya II-III, 376; 
Govorukhin. Fl. Urala 520; Kryl. Fl. Zap. Sib. XI, 3028; Klok. in Vizn. 
Rosl. URSR 603; Szafer, Kulcz., B. Pawl. Rogl. Polskie (1953) 744; Kirp. 
in Majevski, Fl. (1954) 633; Hayek in Hegi, 111, Fl. 2 Aufl. VI, 2, 1118; 
Dostal, Klic (1954) 801; Stank, and Tal. Opred. Vyssh. Rast. (1957) 442; 
Mikhailovskaja in Fl. BSSR, V, 192; Askerova in Fl. Azerb. VIII, 556; 
Kovalevskaja in Fl. Uzb. VI, 487. — L. scariola [3. hortensis 8. sativa Bisch. 
Beitr., z. Fl. Deutschl., Cichorieen (1851) 190. — L scariola p. sativa 
300 Moris, Fl. Sard. II (1840-1843) 531.— L. scariola (3. sativa Boiss. Fl. or. 
Ill (1875) 809.— Ic: Rchb. f. Ic. fl. Germ, et Helvet. XIX (1859) t. 70 
(MCCCCXXI) fig. 7-14; Javorka and Csapody, Iconogr. fl. Hungar. (1933) 
t. 561, fig. 4029; Syreistsch. 111. Fl. Mosk. Gub. Ill (1910) 331; Hegi. op. 
cit. 1119, fig. 797. 

Annual or biennial. Stem 30-70(100) cm high, smoodi, more or less 
sulcate. Leaves gradually strongly reducing upward, almost horizontal, 
lower leaves large, rotund-obovate or elliptical, narrowed into broad 
petiole, sometimes flexuous and often spinose-toothed; middle cauline 
leaves sessile, with deeply cordate base; upper cauline leaves variable, 
often orbicular-reniform, with auriculate or sagittate-amplexicaul base; 
leaves on peduncles very small, almost scaly, densely pubescent. Capitula 
in corymbose-paniculate inflorescence, more or less cylindrical, (6)10- 
13(15) mm long, with 15 florets. Involucral bracts not curved on fruiting, 
three- or four-rowed outer bracts ovate-lanceolate, inner oblong-linear, 
subobtuse, with narrow light-colored border. Corolla yellow. Achenes 
narrow or oblong-obovoid, about 4 mm long and 0.8-1 mm wide, gray 
or (less often) dark brown or brownish, flattened, not always strictly 
symmetrical, with large number of longitudinal ribs on each side, usually 
with erect hairs in upper part (under a lens!), rest smooth, apically attenuate 
into filiform, whitish or light brown beak, almost as long as achene and 
terminating in broad disk bearing pappus; pappus hairs numerous, thin, 
white, soft, slightly barbed (under a hand lens!), almost as long as achene. 
Flowering (May) July to August. (Plate XVIII, Fig. 3.) 

Widely, though not commonly, cultivated in kitchen gardens as a 
vegetable for salads; sometimes found in wild. 

Note. The native country of L. sativa is unknown. This species has 
been considered by many authors as a cultivated variety of L. serriola. 
However, L. sativa is distinguished from the latter by the shape and 
arrangement of the leaves (almost horizontal and not vertical), nature of 
the involucral bracts (after the maturation of fruits, they are deflexed in 



298 

L. serriola, and straight in L. sativa, and also, usually, by the broad 
inflorescence, whose branches are densely covered with small, sessile 
leaves, which are reduced upward; the many capitula in the inflorescence 
often do not develop fully and become globose. The latest information 
on the origin of cultivated lettuce can be seen in Lindquist "On the origin 
of cultivated lettuce." Hereditas, Vol. XL VI, 3^, 1960). 

Economic Importance. The cultivated varieties of L. sativa are 
annuals, which can be cultivated successfully from the extreme south to 
the extreme north both in open fields and under cover (in glass houses and 
hot beds). Lettuce leaves are rich in provitamin A and contain vitamins B,, 
301 B^, C and P and also vitamin E (according to some reports); they are also 
fairly rich in calcium and iron salts; lastly, they also contain phosphorus. 
Such a chemical composition makes lettuce leaves extremely valuable (in 
particular in the diet). Flowering plants are used in homeopathy. 

Information on agronomy and the range of varieties of lettuce can be 
found in V.I. Edelshtein, "Ovshchevodstvo" [Vegetable Cultivation], 2nd 
ed., Sel'khozgiz, Moscow, 1953; and D.D. Brezhnev (Editor)^ "Sorta 
ovoshchnykh kultur" [Varieties of Vegetable Crops], Sel'khozgiz, 
Moscow-Leningrad, 1960. 

The current status of the classification of cultivated lettuce is set 
forth in the book: Mansfeld, Vorlaufiges Verzeichnis landwirtschaftlich 
Oder gartnerisch kultivierter Pflanzenarten, Berlin, Akad.-Verl.,1959. In a 
somewhat simplified form and with small corrections, the infraspecific 
classification of L. sativa can be presented as follows. 

I. Convar. sativa.— Cfr. Helm, Kulturpfl. 2 (1954) 101, 125. 

1. Var. sativa. — L. sativa L.s. str. (quoad typum, L. excl. p. and y. 
Not cultivated. 

2. Var. angustana Irish ex Bremer in Handb. Pflanzenzucht. 5 (1949) 
339 (non Vidi!). — L sativa var. intergrifolia Irish ex Stank, in 
Stank, and Tal. Opred. Vyssh. Rast. (1949) 720. 

This includes the so-called asparagus varieties, in which the 
strongly thickened stem is eaten; leaves narrow, elongated. 

3. Var. longifolia. Lam. Encycl. meth. Ill (1791-1792) 402 (sub nom.: 
y. Lactuca sativa longifolia") 

This includes the lettuce varieties grouped under the name 
"romaine" (in practice, known by the Latin name L. sativa var. 
romana hort."). Their elongated-obovate leaves form lax oblong- 
oval heads. 

II. Convar. incocta Helm, Kulturpfl. 2 (1954) 102, 125. 

4. Var. crispa L. I.e. (sub nom. L. sativa Y. crispa L.). 



299 

It unites the leafy forms not producing heads. In the Russian literature, 
varieties of this group are usually cited under the names, L. sativa var. 
acephala Alef. and L. sativa var. secalina Alef., although Alefeld proposed 
a much more complex structure and such simplified representation of the 
classification — actually quite unwieldy — does not give a correct idea of 
it (cf. F. Alefeld. Landwirtschaftliche Flora ... Berlin, 1866). 

5. Var. capitala L. 1. c. (sub nom. L. sativa p. capitata L.). 

Leaves rotund, forming head. 

The L. sativa x L. serriola hybrids are fertile. This property is used 
for varietal improvement of cultivated lettuce. 

12. L. saligna L. Sp. pi. (1753) 796; DC. Prodr. VII, 1, 136; Ldb. 
Fl. Ross. II, 2, 804, p.p.; Schmalh. Fl. II, 168; Fedtsch. and Fler. Fl. 
Evrop. Ross. 1058; Grossh. Fl. Kavk. IV (1934) 257 and Opred. Rast. 
302 Kavk. 516, p. p.; Nevski in Somye Rast. SSSR IV, 343; Kraschen. in Fl. 
Yugo-Vost. VI. 470; Klok. in Vizn. Ro<?l. URSR, 604; Schchian in R. 
Gruzii VIII, 691; Szafer, Kulcz., B. Pawl. Rogl. Polskie (1953) 743; Hayek 
in Hegi, III. Fl. 2 Aufl. VI, 2, 1125; Dostal, Klec (1954) 801; Kirp. in 
Majevski, Fl. (1954) 632. Stank, and Tal. Opred. Vyssh. Rast. (1957) 
442; Askerova in Fl. Azerb. VllI, 554, p. p. — L. angustifolia Gilib. Fl. 
Lithuan. Ill (1782) 234; Ej.' Exerc. phytol. I (1792) 188.— L. virosa Habl. 
Fiz. Opis. Tavrich. Obi. (1785) 160 (non L. virosa L.). — L caucasica C. 
Koch in Linnaea XVII (1843) 275 (incl.a. major C. Koch and p. minor 
C. Koch. op. cit. p. 276); Ldb. Fl. Ross. II, 2, 806.— L. cyanea C. Koch 
in Linnaea XXIII (1850) 671.— Ic: Rchb. f. Ic. fl. Germ, et Helvet. XIX 
(1859) t. 69 (MCCCCXX) fig. 1-2; Kraschen. op. cit. 471, Fig. 722; 
Hegi, op. cit. 1126, fig. 803; Dostal, op. cit. 799, fig 2219.— Exs.: Sch. 
Bip., Cichoriaceotheca, Suppl. II, No. 153; Billot Fl. Gall, et Germ. exs. 
No. 1700; Kickxia Belgica, Cent. I, 1865, No. 71; Fl.' exs. reip. Bohem. 
Slov. No. 635. 

Annual or biennial. Roots more or less slender, woody, (3)5-10(15) 
cm long. Stem (10)30-80(100) cm high, rather numerous from base, more 
or less arcuate, or stems few to solitary, erect, weakly sulcate, light 
yellowish, stiff, often virgate from base, solid or almost so. Leaves highly 
variable, green or glaucous, rather stiff, sessile, with sagittate 
semiamplexicaul base, densely but very finely appressed hairy (under a 
hand lens!), sometimes covered with dense, often uncate hairs (bristles) 
beneath along midrib; lower leaves runcinate-pinnatisect or emarginate- 
pinnatipartite, with segments or lobes usually retrorse, with convolute 
margins, covered with few spinules; middle cauline leaves lanceolate- 
linear or linear, entire or somewhat toothed-spinose; upper leaves strongly 
reduced; sometimes all leaves linear. Capitula with 9-15 florets, almost 
cyUndrical at flowering, broadened upward at fruiting, about (10)12-14(18) 



300 

mm long, on slender, virgate, branches in lax spicate raceme; peduncles 
slender, short or strongly reduced, and then capitula subsessile. Involucre 
usually rectangular; involucral bracts greenish or more or less violet, short- 
appressed-hairy (under a hand lens!) on dorsal surface; inner bracts 
membranous-bordered, with convolute margins after fruiting, more or less 
strongly divaricate or deflexed. Corolla yellow, blue on drying. Achenes 
about 2.5 mm long and 0.6-0.7 mm wide, narrow-obovoid or oblong- 
ellipsoid, flattened, dark brown or olive-colored, usually with numerous 
prominent longitudinal ribs covered with stiff, upward-directed, very short 
hairs, more distinct in upper part of achene (under a hand lens!), achenes 
apically attentuate into short collar, with whitish filiform beak, usually 
one and one-half to two times as long as achene, terminating in small 
303 funnel-shaped disk bearing pappus; pappus of fragile, very thin, white, 
silky, apparently smooth (under high magnification, weakly barbed) hairs, 
about 5 nmi long. Flowering (June)July to August(September). (Plate 
XVIII, Fig. 8.) 

Dry sunny places, on saUne, calcareous and stony soils, in steppe 
and semidesert, as a weed along banks of rivers, lakes and canals; usually 
solitary or in small groups. — European Part: Middle Dnieper (southeastern 
part), Volga-Don (southern part). Trans- Volga Region, Bessarabia, Black 
Sea Region, Crimea, Lower Don, Lower Volga; Caucasus: Dagestan (on 
lowlands). Eastern and Western Transcaucasia. General distribution: 
central Europe, Atlantic Europe, Mediterranean, Balkans- Asia Minor. 
Described from France and Leipzig. Type in London. 

Note. The general distribution given is approximate (cf. note to L. 
altaica). L. saligna is a highly polymorphic species, from which the 
western European botanists have segregated a series of forms, varieties, 
or even individual races; they also recognize the occurrence of the hybrid: 
L. saligna x L serriola. 

13. L. altaica Fisch. and Mey. in Ind. Sem. Hort. Petrop. XI (1846) 
73; Kryl. Fl. Zap. Sib. XI, 3028; O. and B. Fedtsch. Perech. Rast. Turk. 
IV, 321 (Consp. Fl. Turk. No. 2785); Pavlov, Fl. Tsentr. Kazakhst. Ill, 
365; Popov in Tr. Uzb. Gos. Univ. Nov. Ser. No. 27,' Biol. Vyp. 14, 
103; Kirp. in Fl. Turkm. VII, 341; Vvedensky et al. Opred. Rast. Golodnoi 
Stepi 40 (Tr. Tashk. Gos. Univ. Vyp. 178, Bot.); Kovalevskaja in Fl. 
Uzb. VI, 484. — L. saligna L. var. caule setoso and var. foliis laveibus 
Hohenack. in Bull. Soc. Nat. Mosc. XI, 3 (1838) 270.— L. saligna y. 
robusta Fisch. and Mey. in Ind. Sem. Hort. Petrop. V (1838) 37. — L. 
saligna auct. non L.: Ldb. Fl. Alt. IV, 155; O. and B. Fedtsch. Perech. 
Rast. Turk. IV, 321 (Consp. Fl. Turk.; No. 2784); Kryl. Fl. Zap. Sib. XI, 
3027; Kitam. in Res. Kyoto Univ. Exp. Karak. a. Hinduk. 1955, II, 430 
(Fl. of, Afghanistan, 1960); Zakirov, Fl. i Rastit. Bass. Zeravsh. II, 428.— 
Exs.: Kotschy, PI. alepp. kurd. moss. 1843, No. 442b (sub, L. saligna L. 



301 

var. wallrothii); Sintenis, It. transcasp.-pers. 1900-1901, No. 982b (sub. 
L. saligna L.) 

Annual or biennial. Roots more or less slender, woody. Stem 50- 
80(120) cm high, usually solitary (less often more or less numerous), 
simple or branched, stiff, slightly sulcate, stramineous or whitish, but 
usually violet below and more or less densely spinulose. Leaves glaucous, 
dense, sessile, at base sagittate, glabrous; lower leaves (sometimes partly 
also middle ones) pinnatipartite or erose-lobate, usually with deflexed 
lobes; middle cauline leaves linear, linear-lanceolate, or (less often) oblong, 
entire or shallow spinose-toothed, usually with prominent midrib beneath, 
often densely covered with yellowish spinules (bristles); upper leaves 
strongly reduced; sometimes all leaves linear-lanceolate. Capitula with 
304 (7)9-15(17) florets, more or less cylindrical at anthesis, later often 
narrowed near middle, broadened above at fruiting, about (10)12-13(16) 
mm long, on slender branches, forming lax racemose-paniculate 
inflorescence. Involucre usually four-rowed; involucral bracts greenish or 
with violet spots; outer bracts appressed hairy on dorsal surface (under a 
hand lens!); inner ones with membranous border, more or less strongly 
divaricate after fruiting, or deflexed down. Corolla yellow, blue or pinkish- 
violet on drying. Achenes 3-4 nrun long and 0.6-0.8(0.9) mm wide, 
narrow-obovoid or oblong-ellipsoid, flattened, brown, dark brown, or 
grayish, with (5)7-9(10), prominent, longitudinal ribs on each side covered 
with light colored, upward-directed hairs (under a lens!), which are 
relatively long, squarrose, and soft in upper part. Achenes covered with 
very short transversely sinuate stripes formed by extremely short hairs 
(under high magnification), apically attenuate into collar, bearing whitish 
or yellowish filiform beak more or less as long as achene, sometimes 
distinctly longer; beak with a disk bearing pappus of fragile, very thin, 
white, silky, more or less smooth, about 5 mm-long hairs (under high 
magnification, somewhat barbed). Flowering June to September. (Plate 
XVIII, Fig. 1.). 

Steppe, semidesert and desert regions on saline lands, stony and 
gravelly river valleys, less often in fallows, and roadsides, and also a 
weed in irrigated fields; in mountains up to 2,000 m. — European Part: 
Trans- Volga Region, Lower Volga (Astrakhan); Caucasus: Dagestan (in 
mountains), Eastern and Southern Transcaucasia, Talysh; Western Siberia: 
Upper Tobol, Irtysh, Altai; Soviet Central Asia: Aralo-Caspian Region, 
Balkhash, Dzhungaria-Tarbagatai (only Dzhungaria?), ?Kyzyl-Kum, 
mountainous Turkmenia, Syr- Darya, Pamiro-Alai, Tien Shan. General 
distribution: eastern Mediterranean, Armenia and Kurdistan, Iran. 
Described from Altai. Lectotype in Leningrad. 

Note. The range of L. altaica needs to be confirmed, especially its 
distribution outside the USSR, where this species passes as L. saligna. L. 
altaica is a polymorphic species, which is related to its wide distribution 



302 




305 Plate XIX. 

1 — Steptorhanphus czerepanovii Kiq)., upper part of plant, root, achene; 2 — S. crassicaulis 

(Trautv.) Kirp., achene; 3 — 5. crambifolius Bge., achene; 4 — 5. persicus (Boiss.) O. and B. 

Fedtsch., achene; 5 — 5. tuberosus (Jacq.) Grossh., achene; 6 — S. petraeus (Fisch. and Mey.) 

Grossh., achene; 7 — S. linczevskii Kirp., achene; 8 — Reichardia dichotoma (Vahl) Freyn.: 

a) outer achene; b) inner achene. 



303 

and semi-weedy mode of life. In particular, specimens from the Caucasus 
differ from the typical and Mediterranean specimens, but the differences 
in the material available to me do not give in any way a definite picture. 
The situation is complicated by the fact that L. altaica evidently forms 
306 hybrids with L. saligna. In some regions of the European USSR, not 
indicated by me for L. altaica, specimens are found which are intermediate 
between this species and L. saligna but distinctly closer to L. altaica. 
These include one specimen from the Zilair District in the Bashkirian 
ASSR, one from the Pavlovsk District in the Voronezh Region, and two 
sheets each from the Derkul steppe and Chistyakovo (Donetsk Region) 

It has been established long since that L. altaica is difficult to 
distinguish from L. saligna. In their original description Fisher and Meyer 
reported that the beak of L. altaica is more or less equal in length to the 
achene, whereas in L. saligna it is one and one-half to two times longer 
than the achene. This character has been given prominence by all 
subsequent authors, although, as a matter of fact, it by no means can 
always serve as the prime character, because in practice the achenes of L. 
saligna and L. altaica rarely differ in their beaks. The color of the achene 
is much more stable (lighter-colored in L. altaica), but even better are 
the long, squarrose, soft hairs on the upper part of the achene in L. altaica. 
Moreover, the lower part of the stem in L. altaica usually is violet and 
almost always covered with spinules, which are quite distinct on the midrib 
at least on some leaves, and lanceolate or linear-lanceolate leaves are 
noticeably dominant. Many plants are found, however, in which only by 
taking all of these characters together is it possible to distinguish L. altaica 
from L. saligna with confidence. 

In the opinion of many authors, separating L. altaica from L. serriola 
is no less difficult, as their achenes are practically indistinguishable, and 
the presence of hybrids between them can hardly be doubted. Nevertheless, 
we do not share the skepticism of M.G. Popov, who wrote (op. cit.), 
concerning L. altaica: "Poorly differentiated from the previous one, it is 
better treated as simply a form of that one." Here "previous" means L. 
serriola. In the present case, differences in the vegetative organs, especially 
in the form of the leaves, are of great help. The leaves of L. serriola, 
except the uppermost, are pinnately cut and stiff; in L, altaica, at least, 
the upper ones (and partly also the lower leaves) are entire, small, stiff. 
Besides, L. serriola belongs to the so-called "compass" plants (laminas 
vertical, with the rib facing the sun), which is not typical of, L. altaica. 
However, only specially designed experiments and observations can 
resolve the question of whether L. altaica is a good species. 

14. L. georgica Grossh. in Grossh. and Schischk. Sched. ad. herb. 
PI. or. exsicc. fasc. I-VIII (1924) 8; Ej. H. Kavk. IV (1934) 257 and 



304 

Opred. Rast. Kavk. 516; Schchian in Fl. Gnizii, VIII, 687; Kirp. in Fl. 
Turkm. VII, 337; Askerova in Fl. Azerb. VIII, 554. — Lactuca virosa. 
auct. non L.: O. and B. Fedtsch. Perech. Rast. Turk. IV, 326 (Consp. Fl. 
Turk. No.' 2781).— Ic: Schchian, op. cit. 689, Plate 440.— Exs.: PI. or. 
exs. No. 25 (iso-typus!). 

Perennial (or biennial?). Plants with robust tap root. Stems mostly 
solitary, 75-- 120(1 50) cm high, glabrous or with sparse, stiff, spinules 
307 below, sulcate, fistular, branched above. Leaves stiff, semiamplexicaul, 
elliptical (except upper leaves), ovate or oblong, with prominent network 
of veins beneath, often toothed-spinose; lower leaves almost entire or cut ' 
to various degrees, basally narrowed into broad-winged petiole, covered 
with stiff, about 2 mm long, spinules along midrib beneath; middle leaves 
usually pinnatisect, gradually reduced upward, often with isolated spinules 
beneath along midrib; upper leaves small, variable in form, mostly 
triangular with deeply emarginate base and auriculate appendages. Capitula 
cyhndrical, with fruits 8-11 mm long, with about 15 florets, borne singly 
or in groups of few, almost sessile or, less often on more or less long 
peduncles; inflorescence paniculate or racemose, often robust. Involucre 
usually four-rowed; outer involucral bracts thinly and densely tomentose 
(and also glandular?), often violet; inner ones less densely pubescent, 
with subobtuse, often violet apex, and mostly with distinct midrib. Corolla 
yellow. Achenes dark violet or almost black, somewhat flattened or flat, 
6-7 mm long, broad obovoid part up to 3-3.5 mm long (about 1.5 mm 
wide), abruptly narrowed at apex into narrow, about 1 mm long, collar 
bearing light-colored or dark brownish, filiform, 2-2.5 mm-long beak, 
with prominent ribs (usually seven) on each side covered with numerous 
light-colored stiff, short, upward-directed hairs (under a hand lens!); 
besides, achenes covered with very short, transversely sinuate, stripes of 
short light-colored hairs (under high magnification!); beak terminating in 
small funnel-shaped expansion bearing flat disk with pappus of thin, silky, 
almost smooth, fragile, 3-3.5 mm-long hairs. Flowering July to August. 
(Plate XVIII, Fig. 7.) 

Banks of mountain rivers, rubble slopes and screes. — Caucasus: 
Dagestan, Eastern and Southern Transcaucasia; Soviet Central Asia: 
mountainous Turkmenia. Endemic (?). Described from Georgia. Type in 
Tbilisi, isotype in Leningrad. 

Section 2. Micranthae Boiss. Fl. or. Ill (1875) 804. — Achenes 
narrow-obovoid or oblong-ellipsoid, with one or rarely three longitudinal 
ribs and distinct fiUform beak as long as achene or two to four times as 
long. Corolla usually lilac of pink, 

Lectotype of section: Lactuca glauciifolia Boiss. 



305 

15. L. rosularis Boiss. Fl. or. Ill (1875) 813; kirp. in Fl. Turkm. 
VII, 341. Ic: Kirp. op. cit. 339, Plate LIII. 

Biennial (or perennial?). Root thick, vertical, branched above, 
308 terminating into few stems. Stems divaricately branched from base, 10- 
25 cm high, vv'ith remnants of petioles of dead leaves in lower part and 
slightly sulcate, glaucous surface, very shortly and unevenly pubescent 
(under a hand lens!), distinctly mixed with glandular hairs; fully developed 
leaves arranged in basal rosula or on lowermost part of stem, glaucous, 
oblong-spatulate or oblong-obovate, undivided, with distinct network of 
veins, finely rugose-plicate, densely appressed hairy (under a lens!), shortly 
and unevenly toothed, with dark cartilaginous spinules; upper cauline 
leaves more or less reduced (to scaly), semiamplexicaul, auriculate at 
base. Capitula cyUndrical or oblong-cylindrical, of 10-20(25) florets, with 
fruits 5-7(10) mm long, in lax compound paniculate inflorescence; 
peduncles usually shorter than capitulum, bearing less often sessile, 
relatively large, glandular hairs. Involucral bracts (two) three- or four- 
rowed, reddish-violet, less often green with violet stripes or spots, thinly 
pubescent on outer surface, mixed with glandular hairs; innermost bracts 
less densely pubescent, often greenish, with thin membranous border. 
Corolla? Achenes about 3 mm long and 0.5-0.75 mm wide, dark or 
blackish-brown, narrow-obovoid or elUpsoid, almost entirely smooth (under 
high magnification, very short whitish hairs visible), more or less flat, 
with one distinct central rib, passing through center of ventral or dorsal 
sides continuing into a whitish, light yellow or greenish beak, more or less 
as long as achene and terminating in almost as long pappus of very thin, 
white, somewhat barbed hairs. Flowering May. (Plate XVIII, Fig. 6.) 

On loose saline sands. — Soviet Central Asia: Kara-Kum (western part). 
General distribution: Iran. Described on the basis of Bunge's collections 
from Iran. Type in Leningrad. 

Note. A rare plant for the USSR, so far found only in the Kazandzhik 
District of the Turkmenian SSR. 

16. L. glauciifolia Boiss. FL or. Ill (1875) 813; Popov, in Tr. Uzb. 
Cos. Univ. Nov. Ser. No. 27, Biol, Vyp. 14, 102; Kirp. in Fl. Turkm. 
VII, 341; Kitam. in Res. Kyoto Univ. Exp. Karak. a. Hinduk. 1955, II, 
428 (Fl. of Afghanistan, 1960); Kovalevskaja in Fl. Uzb. VI, 486.— Ic: 
Kovalevskaja, op. cit., Plate XL VIII, Fig. 2.— Exs.: GRF, No. 3500. 

Annual. Stems slender, (2)7-15(20) cm high, divaricately or 
dichotomously branched. Basal leaves usually rosulate, obovate or broadly 
spatulate, narrowed into broad petiole, irregularly toothed, covered beneath 
with rather long, crisped or flexuous, light-colored hairs; sometimes basal 
leaves pinnatisect, coarsely toothed; cauline leaves strongly reduced, often 



306 

309 almost scaly, amplexicaul. Capitula, on the average, with 10-15 florets, 
cyUndrical to oblong- or conical-cylindrical, with fruits about 18 mm long, 
mature capitula slightly narrowed near middle, usually subsessile or on 
short peduncles, less often on long peduncles, in lax dichotomous- 
paniculate inflorescence. Involucral bracts few, usually three- rowed; outer 
bracts with crisped hairs on dorsal surface (like leaves), inner ones narrow 
bordered; involucral bracts more or less squarrose after fruiting. Corolla 
yellow(?), blue on drying. Achenes about 3 mm long and up to 1 mm 
wide, dark brownish-yellow, narrow-obovoid or oblong-ellipsoid, flat, with 
very short transversely hairy stripes (under a hand lens!), and prominent 
rib in center, rugose-toothed upward along lateral ribs (under a hand lens!); 
achenes apically attenuate into filiform white beak two to three times as 
long as achene and terminating in pappus; pappus hairs thin, white, soft, 
somewhat barbed (under a hand lens!), almost as long as achene or slightly 
shorter. Flowering April to May(July). (Plate XVIII, Fig. 9.). 

Gravelly sands and stony slopes, crevices of granite rocks, on 
gypsiferous deposits — everywhere on more or less saline substrates; from 
foothills up to 3,000 m. — Soviet Central Asia: Kyzyl-kum, mountainous 
Turkmenia, Pamiro-Alai, ? Tien Shan (western part). General distribution: 
Iran. Described from Bunge's collections from Iran. Type in Geneva; 
isotype in Leningrad. 

Note. Hybrids with Lactuca undulata Ldb. are possible. The corolla, 
according to Boissier, is light blue, while according to Popov it is yellow. 

17. L. auriculata DC. Prodr. VII, 1 (1838) 140; Boiss. Fl. or. Ill, 
814; O. and B. Fedtsch. Perech. Rast. Turk. IV, 321 (Consp. Fl. Turk. 
No. 2787); Popov in Tr. Uzb. Gos. Univ. Nov. Ser. No. 27, Biol. Vyp. 
14, 103; Zakirov, Fl. i. Rastit. Bass. Zaravsh. II, 429; Kovalevskaja in Fl. 
Uzb. VI, 487. 

Annual. Stem (5)7-30(50) cm high, slightly sulcate, more or less 
densely covered with rather long, pale, narrow-membranous hairs in lower 
part, bifurcate from base or middle. Leaves highly variable, mostly 
(excluding upper) on winged, near base on semiamplexicaul and more or 
less distinctly auriculate-sagittate petioles, thin, green above, glaucous 
beneath and often with thick midrib, covered with sparse hairs (of same 
type as on the stem); lamina lyrately pinnatisect, with relatively large, 
reniform, reniform-ovate or rotund-ovate terminal segment and a few pairs 
of lateral unequal segments; all segments with shallow, roundish, crenate- 
toothed margin, their apices callose, subobtuse; sometimes most leaves 
crowded in lower part of stem, or laminas of lower and middle cauline 
leaves more or less undivided, panduriform, divided only slightly, and 

310 winged structure of petiole less distinct. Capitula of about 15 florets, 
cyhndrical or cylindrical-conical, with fruits (9)10-11(12) mm long, in 



307 

lax corymbose inflorescence; peduncles very slender, usually longer than 
capitula. Involucre mostly four-rowed; involucral bracts greenish, inner 
ones with membranous border, subobtuse and reddish. Corolla lilac-blue. 
Achenes 2-3 mm long and about 0.75 mm wide, narrow-obovoid, flat, 
dark or blackish brown, usually with three prominent longitudinal ribs on 
each side covered entirely with transversely sinuate stripes of extremely 
short light-colored hairs (under high magnification!), and also with 
numerous, very short, upward-directed, stiff hairs, more distinct on latemal 
ribs (under a hand lens!); achenes terminating in roundish notch, from 
which thin pale, 3.5^.5 mm long beak arises; pappus of rather thin, 
white, silky, almost smooth (under high magnification, often finely 
barbed), 3-4 mm long hairs, separated from disk with difficulty. Flowering 
May to August(October). (Plate XVIII, Fig. 4.) 

Semidesert zone and juniper forest zone up to 2,500 m; among stones, 
on slopes, screes, among coastal gravels, herb thickets and scrubs, 
sometimes a weed of roadsides and in crops. — Soviet Central Asia: 
Dzhungaria-Tarbagatai (only one authentic sheet from Dzhungarian Alatau 
seen),?Syr-Darya, Pamiro Alai, Tien Shan. General distribution: Iran, Indo- 
Himalayas. Described from Wallich's collections from Nepal. Type in 
Geneva? 

Note. It is not clear why this species has been omitted in the latest 
manuals of Siro by Kitamura on the flora of Afghanistan and, especially, 
the flora of Nepal. 

18. L. undulata Ldb. Icon. pi. fl. Ross. II (1830) 12, t. 129; Ej. Fl. 
Alt. Iv (1833) 156; DC. Prodr. VII, 1 (1838) 135 (incl. (3. dentata DC); 
Ldb. Fl. Ross. II, 2, 803; Boiss. Fl. or. Ill (1875) 813; O. and B. Fedtsch. 
Perech. Rast. Turk. IV, 321 (Consp. Fl. Turk. No. 2786); Grossh. Fl. 
Kavk. IV (1934) 255 and Opred. Rast. Kavk. 517; Ling in Contrib. Inst. 
Bot. Nat. Acad. Peip. Ill, 4,189; Pavlov, Fl. Tsentr. Kazakhst. Ill, 365; 
Popov in Tr. Uzb. Gos. Univ. Nov. Ser. No. 27, Biol. Vyp. 14, 103; 
Kryl. Fl. Zap. Sib. XI, 3026; Kirp. in Fl. Turk. VII, 340; Nikitina, Mater, 
po Fl. Sev. Ski. Kirg. Ala-too, 111, No. 1130; Kitam. in Res. Kyoto 
Univ. Exp. Karak. a. Hinduk. 1955, II, 431 (Fl. of Afghanistan, 1960); 
Vvedensky et al. Opred. Rast. Golodnoi Stepi 40 (Tr.Tashk. Gos. Univ. 
Vyp. 178; Bot.); Askerova in Fl. Azerb. VIII, 552; Kovalevskaja in Fl. 
Uzb. VI, 488. — L. undulata var. pinnatipartita Trautv. in Tr. Peterb. Bot. 
Sada, I, 1 (1871) 24; L. undulata var. integrifolia O. and B. Fedtsch. in 
Tr. Peterb. Bot. Sada, XXVIII, 1 (1908) 36, nom. nud.— L. undulata var. 
integrifolia C. Winkl. ex O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 
321, nom. nud. — Ic: Ldb. op. cit. (1830) (color., bona); Kovalevskaja, 
311 op. cit. Plate XL VIII, Fig. 3 (solum achenium). — Exs.: Kar. and Kir. PI. 
Soongor. No. 320; Sintenis, It. transcasp.-pers. 1900-1901, No. 1612. 



308 

Annual. Stems (3) 10-30(60) cm high, solitary or few, more or less 
ribbed or sulcate, bifurcate above or from base, usually glabrous below 
(sometimes weakly pubescent above — visible under a hand lens!). Leaves 
almost undivided and sharply toothed to pinnately cut; lower leaves 
petiolate, sometimes rosulate; cauline leaves sessile, auriculate; upper 
leaves small, often undivided or subentire, more or less sinuate in lower 
half; leaves on terminal branches and peduncles inconspicuous. Capitula 
on short peduncles, with 8-12 florets, cylindrical or cylindrical-conical, 
about 20 mm long, narrowed near middle, in paniculate inflorescence. 
Involucral bracts three- or four-rowed; outer bracts densely pubescent, 
inner ones usually with distinct ventation, more or less pubescent, 
subobtuse and almost always with reddish-violet spots. Corolla pinkish- 
violet or light blue. Achenes about 3.5 mm long and 1 mm wide, narrow- 
obovoid, yellowish-dark-brown, more or less flat, with one distinct 
longitudinal rib and transverse rows of sinuate folds, and with relatively 
thick hairs along ribs (distinct under high magnification, especially in the 
upper part of the achene); beak three to four times as long as achene, 
whitish, with pappus of numerous, very thin hairs in upper part, sUghtly 
longer than achene, slightly expanded at base and with two white rod- 
shaped pendent appendages on achene which float in water (as a result, 
beak seems apical on achene). Flowering April to July (August). (Plate 
XVIII, Fig. 5.). 

At altitudes from a few hundred meters to 2,500 m; steppes and 
semideserts, loessy hills and gypsiferous deposits, shallow soils, rubbly 
and stony slopes, gravels, in dry ditches, sometimes as a weed in old 
fields, everywhere on more or less strongly saline soils. — Caucasus: 
Southern Transcaucasia, Talysh; Western Siberia: Upper Tobol, 
Irtysh, Altai, Soviet Central Asia: all regions. General distribution: 
Eastern Mediterranean, Balkans-Asia Minor (Turkey), Iran, Dzhungaria- 
Kashgaria (Kuldzha). Described from Semipalatinsk Region. Type in 
Leningrad. 

Note. It is an extremely polymorphic species. According to the 
word of those who have seen it in nature, in one small area, growing 
side-by-side under apparently similar conditions, are plants 3-5 and 20- 
30 cm high; the sharp difference in growth vigor produces significant 
fluctuations in the nature of the branching, leaf form and other vegetative 
characters. 

According to S. Kovalevskaja (op. cit., p. 489), at the junction of the 
range of L. undulata and L. glauciifolia forms of the first one (pubescent, 
312 with short appendages of the beak) are found which deviate from the 
type and apparently are of hybrid origin. She also reports L. undulata 
from the southeastern European part of the USSR, but I did not see 
specimens from there. 



309 
Species of Uncertain Position 

19. L. spinidens Nevski in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. I, 
4 (1937) 294; id. in Som. Rast. SSSR IV, 343 (descr. ross.); Kirp. in fl. 
Turkm. VII, 336. 

Annual or biennial. Stem 50-130 cm high, whitish, rather thick (to 
7-8 mm at base); more or less sulcate, in lower half covered with 
yellowish, horizontal, up to 5 mm-long spinules. Leaves glaucous-green, 
somewhat coriaceous; lower and middle cauline leaves elliptical or oblong- 
obovate, with rather dense spinules beneath along very prominent midrib, 
from subentire to coarsely and unevenly toothed, with cartilaginous 
spinules along margin, punctate-pitted beneath; upper leaves small, 
appressed to stem, semiamplexicaul, entire, densely pubescent beneath, 
with auriculate or sagittate base. Capitula in long compound racemose- 
paniculate inflorescence, cylindrical or narrow-campanulate, or about 20 
florets and on peduncles mostly longer than capitula, covered with small 
papulose and densely pubescent scaly leaves, sometimes indistinctly 
merging into involucral bracts. Involucral bracts lax, usually four-rowed, 
triangular to lanceolate-linear, on outer surface more or less papulose and 
densely pubescent, obtuse, often with one or few violet spots; inner bracts 
with wide membranous sinuate border. Corolla pale or lemon-yellow. 
Achenes 3-3.2 mm long and 1-1.25 mm wide (dimensions according to 
Nevski), brown or light brown, narrow- or oblong-obovoid, with prominent 
longitudinal ribs (seven to nine on each side)^ covered with very short, 
upward-directed, somewhat stiff hairs (under a hand lens!), more distinct 
in upper part; achene apex attenuate into thin beak, concolorous with 
achene, beak about one and one-half times as long as achene, terminating 
in pappus of thin, white hairs, almost as long as beak. (Plate XVIII, Fig. 
10.) 

In wheat fields and on stony slopes. — Soviet Central Asia: 
mountainous Turkmenia, Pamiro-Alai (Kugitang). Endemic(?). Described 
from Kugitang. Type in Leningrad. 

Note. The description is based entirely on a few specimens without 
fully mature achenes, which does not allow me to determine precisely 
the position of this species in the system of the genus. 

313 Lactuca spinidens is similar to L. aculeata Boiss. and Kotschy ex 

Boiss. However, L. spinidens differs from the latter mainly by the nature 
of the leaves (having punctate pits on the lower surface, which are absent 
in L. acauleata, but then L. acauleata has a more or less conspicuous 
number of erect and stiff or stiffish hairs over the entire lower surface, 
which L. spinidens lacks); the more or less glabrous peduncles (densely 
glandular-hairy in L. acauleata), and other characters. 



310 

Grossheim twice reported L. acauleata from Transcaucasia (Fl. Kavk., 
Izd. 1-e, IV (1934) 257 and Opred. Rast. Kavk. (1949) 516), reliable in 
both cases, with the suggestion that his discovery needs confirmation. In 
the "Flora of Georgia," covering the territory from which this species has 
been reported, the name L. acauleata is missing. In the Herbarium of the 
Botanical Institute, Academy of Sciences of the USSR, there is 
comparatively good material of L. acauleata, including two isotypes 
(Kotschy, Iter cilicicokurdicum 1859, No. 421). None of the specimens 
seen by us from the territory of the USSR can be identified as L. acauleata. 
Thus, this species should be excluded from the flora of the USSR. 

20. L. mira Pavl in Byull. Mosk. Obshch. Isp. Prir. Otd. Biol. Nov. 
Ser. XLII, 2 (1933) 147,— Ic: Pavlov, op. cit. p. 147. 

Perennial. Rhizome slender, creeping, terminating into slender and 
short vanishing stem, reaching 8-15 cm in height including leaves. Leaves 
two or few, on relatively long (usually about 5 cm) and wide, often intort, 
basally thickened petioles with entire or lyrate, oblong-ovate or ovate, 
flat or coriaceous lamina; lamina somewhat rugose, often reddish, finely 
appressed-hairy on both sides, spinose-toothed, often divided at base, 
obtuse or subacute. Capitula usually with 10 florets (8-13), cylindrical, 
with fruits 12-14 mm long, on axillary peduncles arising in groups of 
one or two (less often several); peduncles usually much longer than 
capitulum, with one to several reduced leaves resembling outer involucral 
bracts. Involucre usually three-rowed; involucral bracts dirty dark red, 
very finely pubescent dorsally (under a hand lens!), narrow membranous 
along margin, more or less obtuse; outer bracts sometimes somewhat 
squarrose, ovate or narrow-ovate; inner ones longer, linear- or triangular- 
lanceolate. Corolla pink. Achenes (according to Pavlov) all alike, glabrous, 
ribbed, slightly flattened, weakly narrowed toward both ends, apically 
expanded into a small disk; pappus snow-white, of equal, many-rowed, 
apparently smooth hairs (under high magnification, somewhat and non- 
uniformly barbed). Flowering August. 
314 Upper mountain zone (altitude 3,500 m), on shifting, wet stony screes. 

Soviet Central Asia: Tien Shan (Talass Alatau Range). Endemic. Described 
from the Dzhebogly-Su River. Type in Moscow; isotype in Leningrad. 

Note. A rare and most interesting plant. It is referred to the genus 
Lactuca only conditionally and temporarily until material with mature 
achenes can be studied. It is quite probable that this species ought to be 
referred not to Prenanthes, as proposed by Pavlov, but to the genus Ixeris. 

Doubtful and Deleted Species 

1. L. hybrida M. Pop. ex Zak. in Tr. Uzb. Gos. Univ., Nov. Ser. 



311 

No. 89 (1958) 166, nomen; Zakirov, Fl. i Rastit. Bass. Zeravsh. II (1961) 
428, nomen. 

A plant growing in the "chul" zone on gypsiferous buttes and in the 
"tau" zone on saline stony slopes of terraces has been reported under this 
name. A description is lacking. K.zakirov (1961) notes in parentheses: 
N."? L. undulata x L. glauciifolia." It is mentioned in the Flora of 
Uzbekistan (Vol. VI, 1962). 

2. L. kochiana Beauverd in Bull. Soc. Bot. Geneva, 2 Ser. II (1910) 
115 (seorsum impressum: Contrib. a 1 'etude des Compos. Ill, p. 17); 
Schchian in Fl. Gruzii VIII, 684. — Mulgedium salicifolium C. Koch in 
Linnaea XXIII (1850) 669; Boiss. Fl. or. Ill, S02.—Lactuca salicifolia 
Grossh. Fl. Kavk. IV (1934) 255 and Opred. Rast. Kavk. 516, non L. 
salicifolia Salisb. Prodr. stirp. (1796) 180. 

Perennial. Root? Creeping rhizomes absent. Stem 75-120 cm high, 
erect, weakly branched above. Leaves glaucous, narrow-elliptical or 
lanceolate, shallow runcinate or entire, emarginate or fme-toothed; lower 
leaves short-petiolate, others sessile. Capitula cyUndrical, in racemose- 
paniculate inflorescence; peduncles short, with scaly leaves. Involucral 
bracts reddish, membranous along margin; outer bracts ovate, noticeably 
shorter than inner, ahnost linear bracts. Corolla dark blue. Achenes shortly 
narrowed into beak, brown. Flowering June to August. 

Dry rocky and sandy places in middle and upper mountain zones. — 
Caucasus: Eastern Transcaucasia (Meskhetia). General distribution: 
Armenia and Kurdistan (former Olty District). Described from Daik 
(Turkey). Type was in Berlin and probably was lost. 

Note. The Herbarium of the Botanical Institute, Academy of Sciences 
of the USSR, dqes not have the authentic material of this species. The 
description given here is based only on the literature.. No botanist has so 
far seen fully mature achenes, which prevents this species from having a 
firm position in the classification. C. Koch referred his own species to 
true Mulgedium, but compared it with the present Cicerbita prenanthoides. 
Careful new collections are urgently needed. 

315 3. L. lipskyi Schischk. ex Grossh. Opred. Rast. Kavk. (1949) 516 

(diagn. brev. ross.) 

It is not a vahdly described species, which must be mentioned because 
this name has been used in the widely known manual of A.A. Grossheim. 
The plant is said to be endemic to the Cherkessia region, where it is 
found along the sea coast and, according to Grossheim, has the following 
characters: biennial, up to 60 cm high, strongly branched, branches long, 
virgate; lower leaves deeply divided into narrow, oblong lobes, upper 
leaves strongly decurrent; capitula with 8-12 florets; corolla yellow; beak 
of achene black. 



312 

I was unable to see the authentic material, but judging from the 
specimens studied, the Black Sea Coast of western Transcaucasia does 
not hava a unique species of its own. Most probably, specimens of L. 
chaixii deviating somewhat from the norm were taken for L. lipskyi. 

4. L. perennis L. Sp. pi. (1753) 796. 

This plant, described from Europe, was reported for Russia by 
Ledebour (R. Ross. II, 2 (1845-1846) 802) for his "Central Russia" region, 
in particular: "Livonia: in montosis p. Wanden (Fisch.) Warsawia (Emdt)." 
Subsequently, Schmalhausen (Fl. Sredn. i. Yuzhn. Ross. II (1897) 167) 
wrote that L. perennis is "reported from the Minsk Province." However, 
Fedtschenko and Flerow (Fl. Evrop. Ross. (1910) 1057) noted that the 
reported occurrence of this species "needs confirmation." Stankov and 
Taliev in their book (Opred. Vyssh. Rast., 2nd ed. (1957) 442) again 
mentioned L. perennis for the Minsk Region, but in the Flora. BSSR 
(Vol. V, 1959) Mikhailovskaja did not mention this species.There is no 
material in the Herbarium of the Botanical Institute, Academy of Sciences 
of the USSR that could be identified with L. perennis. L. 

5. L. quercina L. Sp. pi. (1753) 795; Ldb. Fl. Ross. II, 2, 806; 
Schmalh. Fl. II (1897) 168 (incl. var. sagittata, striata and armena) 
Fedtsch. and Fler. Fl. Evrop. Ross. 1059. 

I could not establish the identity of L. quercina L. This species was 
described by Linnaeus from an island in the Gulf of Bothnia ("Habitat in 
insula Carolina Balthici"), which now carries the Finnish name Hailuoto, 
and earlier carried the Swedish name of Karlo Island). 

DeCandolle {Prodr. VH, 1 (1838) 138, No. 36) aUuded to the doubtful 
status of this species ("Planta... forte dubia"). "Index Kewensis" indicates 
that L. quercina L. is found in southern Europe and the Caucasus ("Europe 
austr., Reg. Cauc"), i.e., in regions having no relationship to the locus 
classicus. On the other hand, assuming that a unique Lactuca species 
exists in the Gulf of Bothnia, it can hardly be expected to reappear, after 
a large gap, in southern European USSR and the Caucasus. However, it 
must be mentioned for the sake of objectivity that western floristic 
316 botanists usually recognize L quercina, and, besides, most treat this species 
very broadly, including L chaixii and L, stricta as subspecies (or varieties). 

Possibly, Linnaeus described L. quercina from the extreme northern 
outpost for the species, and then at least L. stricta Waldst. and Kit. 
becomes a synonym of L. quercina L. 

6. L. querna Pall. Bemerk. Reise in die Sudl. Statth. d. Russ. Reichs 
I (1799) 221, solum nomen. 

In section "Astrachanische Weinbau," Pallas opens the list of 



313 

commonly occurring plants with "wild Salat (Lactuca quema)." Ledebour 
ignored this name of Pallas. Trautvetter (Tr. Peterb. Bot. Sada VIII, 2, 
1883, Increm. fl. phaenog. Ross. fasc. II, p. 521, Nov. 3247) quite 
categorically states that the plant of Pallas is nothing other than L. quercina 
L. In Index Kewensis, this same opinion was also expressed, with a 
question mark. The real difficulty is that L. quercina, as understood by 
the older Russian authors from the Astrakhan area, is as yet not known. 
Most probably, Pallas saw a wild specimen of cultivated lettuce, but this 
question can be resolved finally only after examining the authentic 
specimen. 

7. L. spectabilis Fisch. and Mey. ex. Sch. Bip. in Herb, et Winkl. 
Jahrb. Pharm. IV (1841) 154, in nota (vidi tantum seorsum impressum 
sub titulo "Sceleton systematis articulati Cichoriacearum," p. 4). 

Schultz, making the new combination Wiestia spectabilis (Fisch. and 
Mey.) Sch. Bip., took Lactuca spectabilis Fisch. and Mey. as the basionym. 
It can be established by indirect information that a plant of L. serriola 
was considered under this name. So far I have not been able to establish 
precisely what L. spectabilis is and from where it was described. The 
name L. spectabilis is not mentioned in "Index Kewensis." 

8. L. tephrocarpa C. Koch in Linnaea XXIII (1850) 672. 

This species was described by C. Koch from Georgia. However, it 
has not been mentioned subsequently by Russian investigators of the flora 
of Caucasus (Lipsky, Grossheim and others). Probably, one has to agree 
with Boissier (who had seen the material of Koch) and the authors of 
"Index Kewensis" that L. tephrocarpa is not different from L. serriola. 

9. L. virosa L. Sp. pi. (1753) 795 and ed. 2 (1763) 1119; DC. Prodr. 
VII, 1,137; Ldb. Fl. Ross. II, 2, 805; Schmalh. R. II, 167; Fedtsch. and 
Fler. Fl. Evrop. Ross. 1058; Kraschen. in R. Yugo-Vost. VI, 470; Szafer, 
Kulcz., B. Pawl. Rogl. Polskie (1953) 743; Kirp. in Majevski, Fl. (1954) 
633; Hayek in Hegi, 111. Fl. 2 Aufl.VI, 2, 1122; Dostal,KJec (1954) 801; 
Stank, and Tal. Opred. Vyssh. Rast. (1957) 443. — ^Mikhailovskaja in Fl. 

317 BSSR, V, \9\.— Wiestia virosa Sch. Bip. in Herb, et WInkl. Jahrb. Pharm. 
IV (1841) 154, in nota (vidi tantum seorsum impressum sub titulo 
"Sceleton systematis articulati Cichoriacearum", p. 4). — Ic: Hegi, op. cit. 
1123, fig. 800.— Exs.: Sch. Bip. Cichoriaceotheca, No. 82; Kickxia 
Belgica, No. 329. 

Ledebour cited this species only on the basis of the published reports 
of Falk (Volga below Kazan), Georgi (Saratov), and Uspensky 
(Sverdlovsk). Ledebour did not see specimens ("Specimina rossica nodum 
vidi"). Schmalhausen indicates an apparently rather wide distribution of 



314 

this species (Lithuania, Belorussia, Ukraine, and even Turkestan!). This 
opinion of Schmalhausen, based on imprecise identifications, was repeated 
by Fedtschenko and Flerow and, among the later authors, Stankov. Other 
Soviet taxonomists (Krascheninnikov, Mikhailovskaja and others) generally 
cited this species with various cautions, emphasizing the need to confirm 
the locality of L. virosa or indicating that they did not see herbarium 
specimens. 

All specimens, that we were able to examine in the Herbarium of the 
Botanical Institute, Academy of Sciences of the USSR, with the name L. 
virosa L. were simply wrongly identified plants (most of these belong to 
L. serriola). Thus, at present there is no basis to consider L. virosa as 
growing within the territory of the USSR. 



GENUS 1655. ScariolaF.W. Schmidt'-^ 

F.W. Schmidt, Samml. phys.-okonom. Aufs. I (1795) 270 (non vidi!); 
Sojak in Novit. bot. Horti bot. Univ. Car. Prag. (1962) 41, 46. — 
Phaenixopus Cass, in Diet. sc. nat. XXXIX (1826) 391. — Phaenixopus 
Rchb. Consp. (1828) 9S.—Phaenopus DC. Prodr. VII, 1 (1838) 176.— 
Lactuca sect. 2. Phaenixopus Benth in Benth and Hook. f. Gen. PI. II 
(1873) 525, p. p. — Lactuca sect. Quinqueflorae Boiss. Fl. or. Ill (1875) 
805, p. p.—Phaenopus DC. Num. Consp. Fl. Europ. II (1879) 435.— 
Lactuca subgen. Phaenixopus Babcock, Stebbins and Jenkins in Cytologia, 
Fujii Jubil. Vol. (1937) 191, solum, nomen. 

Capitula homogamous, mostly five-flowered, cylindrical, borne in 
spicate-panicles or spicate inflorescence. Involucre three- or four-rowed; 
bracts herbaceous, innermost membranous along margin, subobtuse. 
Corolla yellow. Achenes mostly flattened, pubescent with more or less 
numerous (usually five to seven or seven to eight) prominent longitudinal 
ribs (under a hand lens!), apically attenuate into long beak-like structure, 
or only in narrow throat-like structure sometimes achenes narrowed almost 
equally on both sides, narrow-ellipsoid, with distinct or indistinct 
318 narrowing at apex. Pappus of numerous, fine, fragile haris. Basic 
chromosome number in cytologically studied species is jc = 9. 

Type of genus: Scariola viminea (1.) F.W. Schmidt. 

Biennial and perennial herbs or semishrubs, usually growing in arid 
regions, often on stony places. 



' Treatment by M.E., Kirpicznikov. 

~ A new Latin word derived from the local Italian name scariola, used for salad 
plants, including also species of the genus Lactuca s. 1. 



315 

The genus comprises about 10 species, growing in the Old Worid: in 
the USSR mainly in the Caucasus and Soviet Central Asia, as also in the 
southern half of the European Part; outside the USSR, in Europe 
(especially in the Mediterranean countries), Asia Minor, Southwest Asia 
(Turkey, Syria, Lebanon, Iran etc.), Afghanistan and, possibly, the 
Himalayas and Tibet. 

In the structure of achene, Scariola species are close to the members 
of subgenus Lactucopsis of the genus Lactuca, as its narrowed apex, even 
if appearing like a beak, is not structurally separated and is simply the 
apical part of the achene. However, the structure of the vegetative organs, 
ecology and also the few-flowered capitula (with practically a fixed 
number of florets), clearly distinguish the Scariola species from 
Lactucopsis. 

1. Achenes dark violet or blackish, apically attenuate into long beak- 
like part. Biennial or perennial plants 

1. S. viminea (L.) F.W. Schmidt. 

+ Achenes brown- dark brown or yellowish 2. 

2. Achenes 7-8 mm long 2. S. orientalis (Boiss.) Sojak 

+ Achenes 2.5-3 mm long 

3. S. albertoregelia (C. Winkl.) Kirp. 

1. S. viminea (L.) F.W. Schmidt, Samml. phys.- okonom. Aufs. I 
(1795) llO.—Prenanthes viminea L. Sp. pi. (1753) 797; M.B. PL taur.- 
cauc. II, 244. — Phoenixopus vimineus Rchb. Fl. Germ, excurs. II (1831) 
272; Sosnowsk. in Takht. and Fed. Fl. Erevana (1945) 339 (sub 
Phaenixopode vimineo). — Prenanthes ramosissima All Fl. Pedem. I (1785) 
226. — Chondrilla sessiliflora Lam. Fl. Fr. II (1778) 104. — C. viminea 
Lam. Encycl. meth. II (1786) 77. — Phaenixopus decurrens Cass, in Diet. 
sc. nat. XXXIX (1826) 391, p. ^.—Lactuca viminea J. and C. Presl. Fl. 
Cech. (1819) 160; Schmalh. Fl. II, 169; Fedtsch. and Fler. Fl. Evrop. 
Ross. Fl. 1058; Grossh. Fl. Kavk. IV (1934) 256 and Opred. Rast. Kavk. 
516; Kraschen. in Fl. Yugo-Vost. VI, 470; Klok. in Vizn. Rosl. URSR, 
604; Schchian in Fl. Gruzil VIII, 691; Hegi, 111. Fl. 2 Aufl. VI, 2, 1129; 
Dostal, Klec. 802; Kirp. in Majevski, Fl. (1954) 632; Stank, and Tal. 
Opred. Vyssh. Rast. (1957) 442; Askerova in Fl. Azerb. VIII, 556.— 
Phaenopus vimineus DC. Prodr. VII, 1 (1838) 176. — Lactuca viminea 
Sch. Bip. ex. W.D. Koch Taschenb. d. Deutsch. u. Schweiz. Fl. (1844) 
317; Ldb. Fl. Ross. II, 2, 804.— Ic: All. op. cit. t. XXXIII, fig. 1 (sub 
Prenanthe ramosissimo); Rchb. f. Ic. fl. Germ,, et Helvet. XIX (1859), 
319 t. 67 (MCCCCXVIII), fig. 1-9 (sub Lactuca viminea ut omnia sequen.); 
Javorka and Csapody, Iconogr. fl. Hungar. (1933) t. 561, fig. 4025 (bona); 
Hegi, op. cit. 1129; fig. 806 (achenium non exacte delineatum); Dostal, 



316 

op. cit. 803, fig. 2221.— Exs.: Rchb. Fl. Germ. exs. No. 598 (sub 
Phaenixopode vimineo); Sintenis, It. trojanum 1883, No. 476; Ej. It. 
transcasp.-pers. 1900-1901, No.l002; Ross, Herb. Siculum, No. 554 (tria 
ultima sub Lactuca viminea). 

Biennial (or perennial?). Root woody, long, more or less vertical. 
Stem 30-60(80) cm high, glabrous, light colored (straw-yeUow or whitish), 
branched from middle or base, with long and slender, virgate branches, 
less often stem simple. Leaves, especially upper ones, finely tomentose 
(under a lens!); basal and lower cauline leaves early-withering, pinnatipartite 
or pinnatisect, with lobes or segments usually linear-lanceolate, somewhat 
toothed or entire; other leaves sessile, usually linear, entire, decurrent, 
with long, linear bases (auricles) extending downward, distinguished by 
their green stripes on light-colored stem. Capitula with five florets, 
cylindrical, or almost cylindrical, with fruits up to 22 mm long, in groups 
of few or solitary, in spicate-panicles or spicate inflorescence, mostly 
sessile; terminal capitula often on long peduncles, with scaly leaves. 
Involucre usually four-rowed, involucral bracts herbaceous, greenish, finely 
appressed hairy on dorsal surface (under a hand lens!), from short, 
triangular-ovate outer bracts to longer and linear inner with membranous 
border, subobtuse. Corolla yellow. Achenes flattened, with rather numerous 
(usually seven to nine), prominent longitudinal ribs, dark violet or blackish, 
covered with numerous short and stiff hairs (under high magnification!); 
lower, relatively broad, ellipsoidal part of achenes about 5 mm long and 
up to 1 mm wide, gradually attenuate upward, forming beak-like structure 
bearing light-colored disk with pappus; total length of achene with beak- 
like structure 12-14 mm; pappus of numerous white, fine, silky, somewhat 
barbed (under a hand lens!), fragile, 5-6 mm long hairs. 2n = 18. Flowering 
June to September. (Plate XXIII, Fig. 2.). 

Forest-steppe and steppe regions in stony and weedy places; in lower 
and middle mountain zones on stony and rubbly slopes and on screes. — 
European Part: Volga-Kama (Tatar ASSR), Trans-Volga Region, 
Bessarabia, Black Sea Region, Crimea, Lower Don; Caucasus: Ciscaucasia, 
Eastern, Western, and Southern Transcaucasia, Talysh; Soviet Central Asia: 
mountainous Turkmenia (less often). General distribution: central Europe, 
Atlantic Europe, Mediterranean, Balkans-Asia Minor, Armenia and 
Kurdistan, Iran. Described from France and Portugal. Type in London. 

Note. The information about its distribution in the European USSR, 
except for the territory of Crimea, comes only from published sources. 
320 Localities in Soviet Central Asia as well as outside the USSR need to be 
confirmed. 

2. S. orientalis (Boiss.) Sojak in Novit. bot. Horti bot. Univ. Car. 
Prag. (1962) 46. — Phaenopus orientalis Boiss. Voy. Bot. Esp. II (1839- 



317 

1845) 390, in nota; Nevski in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 4 
(1937) 294; Zakirov, Fl. i Rastit. Bass. Zeravsh. II, 427. — Phaenixopus 
decurrens Cass, in Diet. sc. nat. XXXIX (1826) 391, p. min. p. — Lactuca 
viminea Sch. Bip. p. erostris Rgl. in Bull. Soc. Nat. Mosc. XL, II, 3 
(1867) 173., — L. viminea Sch. Bip. var. leucocarpa Trautv. in Tr. Peterb 
Bot. Sada II, 2 (1873) 559.— L. orientalis Boiss. Fl. or III (1875) 819; O. 
and B. Fedtsch. Perech. Rast. Turk. IV, 332 (Consp. Fl. Turk. No. 2789); 
Grossh. Fl. Kavk. IV (1934) 256 and Opred. Rast. Kavk. 516; Kirp. in 
Fl. Turkm. VII, 438; Kitam. in Res. Kyoto Univ. Exp. Karak. a. Hinduk. 
1955, 430 (Fl. of Afghanistan, 1960); Askerova in Fl. Azerb. VIII, 556; 
Kovalevskaja in Fl. Uzb. VI, 483. — Mulgedium orientale (Boiss.) M. Pop. 
in Tr. Uzbek. Gos. Univ. Nov. Seriya, No. 27, Biol. Vyp. 14 (1941) 
104. — Phoenixopus orientalis Nevski ex M. Pop. ibidem, nom. in synon. — 
Phaenixopus orientalis Boiss. ex. Sosn. in Takht. and Fedtsch. Fl. Erevana 
(1945) 339.— Ic: Bouloumoy, Fl. Lib. et Syr. (1930) t. 226, fig. 4 (phot., 
mediocr., sub Lactuca orientali). — Exs.: Kotschy, PI. Pers. austr. No. 702 
(sub Phaenopode orientali); Bomm. It. pers.-turc. (1892-1893), No. 4120 
(sub Lactuca orientali); Bomm. It. pers.-turc. (1892-1893), No. 4120 (sub 
Lactuca orientali, ut ulteriora); Sintenis, It. or. 1890, No. 2782 and It. or. 
1892, No. 4785; Ej. It.transcasp.-pers. 1900-1901, Nos. 587b, 919a and 
b, 1090; GRF No. 1885. 

Perennial. Semishrub with somewhat strong, woody, fibrous branched 
roots. Plants divaricately branched from base. Stems (10) 20-60 cm high, 
woody below, glabrous or subglabrous; branches stiff, virga^e, sometimes 
sulcate, whitish to snow-white, with green striations of decurrent bases of 
leaves, apical leaves often spinescent. Leaves glaucous-green, rather dense, 
subglabrous (under high magnification, finely appressed-hairy); lower 
leaves (early-withering) pinnatipartite or pinnatisect, with two to four pairs 
of usually triangular lobes or segmens and long, linear, acuminate apex, 
narrowed toward base into broad semiamplexicaul petiole; other leaves 
sessile, decurrent with long, narrowly linear bases (auricles), fused with 
stem; middle cauline leaves with laminas like lower leaves in shape, upper 
linear, entire. Capitula usually with five-florets, narrowly cylindrical, with 
fruits 8-14 mm long, solitary, or, less often, in groups of few, on branches 
in spicate or spicate-paniculate inflorescence, usually sessile or terminal 
on more or less long peduncles covered with scaly leaves. Involucre three- 
or four-rowed; involucral bracts herbaceous, green to (partly or wholly) 
violet; outer bracts ovate, densely fine tomentose on dorsal side, inner 
321 ones much longer, with membranous border, subobtuse. Corolla yellow. 
Achenes 7-8 mm long and 1-1.25 mm wide, narrowly ellipsoid, flattened, 
brown, drark-brown or yellowish, less often dirty dark red, with five to 
seven longitudinal ribs (some not always prominent), covered with very 
short hairs (under a lens!), apically attenuate into short (sometimes 



318 

indistinct), narrow, throat-like structure bearing light-colored disk with 
pappus; pappus of numerous white, fine, silky, somewhat barbed (under 
a hand lens!), fragile hairs, as long as achene or slightly shorter. 2n = 36. 
Flowering June to September(October). (Plate XXIII, Fig. 1.). 

Foothills and mountains up to 3,800 m; on clayey and loamy soils, 
lime deposits and shales, dry rubbly and stony places, mostly on slopes 
and screes, as well as on gravels and in dry ravines. — Caucasus: Southern 
Transcaucasia, Talysh: Soviet Central Asia: Kyzyl-Kum Kara-Kum, 
mountainous Turkmenia, ?Amu-Darya, Syr-Darya, Pamiro-Alai, Tien Shan. 
General distribution: Mediterranean, Balkans-Asia Minor (Asia Minor), 
Armenia and Kurdistan, Iran, Indo-Himalayas (Himalayas; literature 
records), Tibet (literature records). Described from southwest Asia. Type 
in Geneva. 

Note. A highly polymorphic species, but so far not yielding to division 
into well-defmed races. Within Soviet Central Asia, individual specimens 
of 5. orientalis differ markedly from each other both in size and growth 
habit: some plants are relatively tall and lax, while others are almost 
pulvinate and produce forms with numerous, slender, very dense, often 
more or less entangled and spinose branches; these dwarf forms also differ 
by having relatively short capitula and almost violet involucral bracts. 
However, I did not fmd specimens with mature achenes, in the absence 
of which it is difficult to judge the taxonomic significance of such plants. 
Most likely, this is simply an alpine variant of S. orientalis. In a series of 
Mediterranean plants, especially those growing in the Gomyi Badakhshan, 
small pads consisting of clusters of white arachnoid silky hairs are 
observed at the base of the expanded dead petioles of the lower 
leaves. 

However, the plants growing in the western Tien Shan deserve special 
attention. Two sheets preserved in the Herbarium of the Botanical Institute, 
Academy of Sciences of the USSR, have the following label: "Compositae. 
Lactuca pseudorientalis M. Pop. sp. n. Middle reaches of Kok-Su, granite 
scree." This whole label, as Linczevsky kindly informed me, was written 
by O.K. Smirnova, the wife of M.G. Popov. On the same label, in 
Gorshkova's hand, is the postscript: "collected by O.K. Smirnova." These 
specimens differ from others by having a relatively large number of rows 
(five to seven) of greenish involucral bracts (quite distinct in young 
capitula) as well as crowded capitula and almost green branch tips because 
of leaves densely covering their bases. Unfortunately, the absence of 
322 mature achenes does not allow us to express a firm opinion about "L. 
pseudorientalis" but it appears that this is only a form of S. orientalis 
and not a separate race. 

The specimens from Badkhyz (collections of LA. Linczevsky from 
the Gyaz-Gyadyk Range, 1930) also attract attention by their general 



319 

morphology. The secondary branches are particularly strongly branched 
above, the involucral bracts are mostly four- or five-rowed, and leaf bases 
on fully mature specimens are almost unnoticeable. However, the achenes 
are practically indistinguishable from typical S. orientalis. 

The aforesaid speaks to the need for a more detailed study of S. 
orientalis s. 1. in the field; also desired are new collections of complete 
specimens with mature achenes. 

3. S. albertoregelia (C. Winkl.) Kirp. comb. nova. — Chondrilla 
albertoregelia C. Winkl. in Tr. Peterb. Bot. Sada XI, 10 (1890) 337 
(seorsum impressum; in volumino completo XI, 2 (1892) 337); O. and B. 
Fedtsch. Perech. Rast. Turk. IV, 323 (Consp. Fl. Turk. No. 2795).— Crepis 
aff. multicaulis Iljin in Byull. Otdel. Kauchukon. 3 (1930) 61. 

Perennial. ? Plants 20-30 cm high, with dead remnants of 
membranous sheaths of lower leaves at base, mostly covered with 
malpighian or stellate hairs. Stem slender, virgate, branched from base, 
glabrous or subglabrous, sometimes with stellate hairs only in lower part. 
Lower leaves oblong-lanceolate or oblong-linear, narrowed into winged 
petiole, merging with narrow-triangular sheath-like base; cauline leaves 
sessile, narrow-linear to subulate, inconspicuous, scaly at stem apices. 
Capitula with five florets, cylindrical, with fruits 6-7 mm long, solitary 
at apices of slender branches, far exceeding capitula. Involucre (two) three- 
or four-rowed; involucral bracts herbaceous, brownish, inner bracts usually 
five, much longer than outer, with membranous border, subobtuse. Corolla 
yellow(?). Achenes 2.5-3 mm long and 0.3 mm wide, smooth or almost 
so, slightiy flattened or not, brown-dark brown, usually with five prominent 
324 longitudinal ribs, apically attenuate into beak-like structure (about 0.5 
mm long), bearing broad disk with pappus of thin, white, somewhat barbed 
(under a hand lens!), usually fragile hairs, slightly longer than achene. 
Fruiting September. 

Valleys of mountain rivers to altitudes of 2,000-2,500 m. Soviet 
Central Asia: Pamiro-Alai (Darvaz). Endemic. Described from Darvaz. 
Lectotype and paratypes in Leningrad. 

Note. Described on the basis of a small number of authentic 
specimens, among which there are no capitula with florets, and the achenes 
are clearly immature. Therefore, the taxonomic status of this species, 
particularly, the assigning of it to the genus Scariola, remains doubtful. 
However, it is clear that S. albertoregelia cannot remain in the genus 
Chondrilla, to which K. Winkler refers it , nor in Crepis as proposed by 
Iljin. Based on achene structure, the plant examined is close to species of 
Lactuca L. s. 1., including also Scariola. It differs from the typical members 
of the latter genus by its leaf structure and particularly by the nature of 
the hairs on the sheaths of the lower leaves; the hair type (malpighian or 



320 




323 Plate XX 

1 — Cephalorrhynchus kirpicznikovii Grossh., habit of plant, achene; 2 — C. tuberosus (Stev.) 

Schchian, achene; 3 — C. talyschensis Kirp., achene; 4 — C. soongoricus (Rgl.) S. Kovalevsk., 

achene; 5 — C. kossinskyi (Krasch.) Kirp.; 6 — C. subplumosus S. Kovalevsk., achene; 7 — C 

takhtadzhianii (Sosn.) Kirp., achene; 8 — C. polycladus (Boiss.) Kirp., achene; 9 — Mycelis 

muralis (L.) Dum., achene; 10 — Lagoseriopsis popovii (Krasch.) Kirp., achenes: a) outer; b) 



321 

stellate) alone sharply differentiates this species from the other members 
of the genus Lactuca L. s. 1. 

GENUS 1656. Steptorhamphus Bge.'^ 

Bge. in Beitr. z. Kennth. Fl. Russl. u. Stepp. Centr.-As. i.e. Alexandri 
Lehmann reliq. hot. etc. (1851) 205 (seorsum impressum) and in Mem. 
Sav. Etr. Petersb. VII (1854) 381. — Lactuca sect. 4. Cicerbita Benth. in 
Benth. and Hook. f. Gen. PI. II (1873) 526, p. min. p. — Lactuca sect. 
Tuberosae Boiss Fl. or. Ill (1875) 804. — Cicerbita §5. Steptorhamphus 
Beauverd in Bull. Soc. Bot. Geneve, 2 Ser. II (1910) 117 (seorsum 
impressum, Contrib. a I'etude des Compos. Ill, p. 19). 

Capitula homogamous, with (15)25-35(50) florets relatively large 
(with fruits up to 40 mm long), cylindrical, narrow-campanulate or oblong, 
in corymbose, corymbose-paniculate, racemose or racemose-paniculate 
(sometimes as if candelabriform), inflorescence. Involucral bracts 
imbricate, usually three- or four-rowed; innermost bracts two times or 
more as long as outer ones, usually linear, with membranous border. 
Receptacle smooth. Anthers oblong-linear, with rather long, appressed, 
more or less sagittate basal appendages. Style and its branches densely 
covered with pointed, upward-directed, short hairs. Achenes somewhat 
ellipsoid or lanceolate, less often oblong, flattened to completely flat, 
with one or three, more or less distinct, ribs on each side, densely covered 
with very short, transverse, sinuate rows of light-colored hairs (under a 
hand lens!), mixed with isolated, relatively long (white and transparent), 
papilliform hairs perpendicular to achene surface; beak thin, usually 
325 yellowish-green. Pappus persistent, of short outer corona (under a hand 
lens!) and inner rows of white , relatively long, thin, silky, more or less 
smooth hairs (under high magnification, somewhat barbed), outer corona 
of numerous hairs in few or many rows, lemon-colored or golden, or 
only of more or less dense, sometimes distinctly reduced, ring of white 
transparent hairs not always conspicuous (under a lens!). 

Type of genus: Steptorhamphus crambifolius Bge. 

The genus comprises about 10 species of perennials with thick and 
often lignified roots. Plants almost exclusively growing in Soviet Central 
Asia and the Caucasus; one species in Crimea; outside the USSR, found 
in Afghanistan, Pakistan, Iran, Turkey, and in the Mediterranean. 

In the original description, Bunge compared his new genus with 
Lomatolepis Cass. Actually, Lomatolepis is relatively distant from 



Treatment by M.E. Kirpicznikov. 

From the Greek words stepto [sic; strepi] — twisted, and rhamphos — ^beak. 



322 

Steptorhamphus, which is phylogenetically related to Lactuca and 
Cicerbitcu 

1. Beak shorter than achene 2. 

+ Beak as long as achene or longer 3. 

2. Involucral bracts (and also peduncles) covered with glandular hairs 

2. S. crassicaulis (Trautv.) Kirp. 

+ Involucral bracts, at least outer, finely appressed-hairy, but without 

glandular hairs 1. S. crambifolius Bge. 

3(1) Peduncles glandular-hairy; outer corona of pappus consisting of 

more or less numerous, often many-rowed, usually golden or 

lemon-yellow, short hairs (under a hand lens!) 

3. S. persicus (Boiss) O. and B. Fedtsch. 

+ Peduncles without glandular hairs; outer corona of pappus usually 

consisting of incomplete row or of fewer, short, white hairs, not 

always conspicuous (under a hand lens!) 4. 

4. Corolla Ught blue-purple or light blue-lemon-colored; beak fiUform, 
two and one-half to three times as long as achene 

4. S. tuberosus (Jacq.) Grossh. 

-I- Corolla yellow or beak shorter 5. 

5. Middle cauline leaves undivided, with shallow incisions; peduncles 
rather slender (immediately below capitulum, 0.5-1 mm wide). 
Plants so far known only from southern Transcaucasia 

7. S. czerepanovii Kirp. 

+ Middle cauline leaves with lamina more or less divided, often 
runcinate 6. 

6. Basal (lower) and middle cauline leaves runcinate, densely 
tomentose (under a lens!); basal leaves with more or less numerous 

326 hairs beneath along with their thick midrib. Plants of the Caucasus. 

5. S. petraeus (Fisch. and Mey.) Grossh. 

+ Basal (lower) and middle cauline leaves undivided, more or less 
coarsely toothed-spinose, less often pinnatilobate and pinnatipartite 
to almost runcinate, finely appressed hairy (under high 
magnification!). Plants from Turkmenia .... 6. S. linczevskii Kirp. 

Section 1. Steptorhamphus. — Outer corona of pappus usually of 
numerous lemon-colored or golden-yellow hairs in a few or many rows; 
hairs on achene all alike. Projections on the aerole of disk usually 
somewhat flat. 

Type of section: type of genus. 

1. S. crambifolius Bge. in Beitr. z. Kennto. Fl. Russl. u. Stepp. Centr.- 
As. i. e. Alexandri Lehmann reliq. bot. etc. (1851) 205 (seorsum 



323 

impressum) and in Mem. Sav. Etr. Petersb. VII (1854) 381; O. and B. 
Fedtsch. Perech. Rast. Turk. IV, 319 (Consp. Fl. Turk. No. 2777); 
Kovalevskaja in Fl. Uzb. VI, 492. — Lactuca crambifolia Boiss. Fl. or. Ill 
(1875) 806; B. Fedtsch. Rastit. Turk. 765.— L. crambifolia (Bge.) M. 
Pop. in Tr. Uzb. Gos. Univ. Biol. Vyp. 14 (1941) 109 (in appendix).— 
Cicerbita crambifolia Beauverd in Bull. Soc. Bot. Geneve, 2 Ser. II (1910) 
143 (seorsum impressum: Contrib. a I'etude des Compos. Ill, p. 45, No. 
50); Vvedensky et al. Opred. Rast. Golodnoi Stepi 40 (Tr. Tashk. Gos. 
Univ. Vyp. 178, Bot.).— Ic: Kovalevskaja, op. cit. Plate XLIX, Fig. 2. 

Perennial. Root 1-1.5(3) cm thick, sometimes branched. Stems 
(15)30-40(50) cm high, glabrous or almost so, sulcate, fistular, usually 
branched from middle or above producing auxiliary floriferous branches. 
Leaves glaucous, rather dense, often with violet tinge; basal leaves often 
rosulate at stem base, eUiptical or oblong-obovate, usually up to 25-30 
cm long and 4-7 cm wide, undivided to pinnatilobate and pinnatipartite, 
or sinuate on margin, in both cases spinulose along margin, with thick 
midrib expanded downward and merging with still broader petiole; cauline 
leaves much smaller than basal, oblong-elliptical, spatulate or lanceolate, 
usually undivided, less often somewhat incised or divided, on broad 
semiamplexicaul petiole or sessile; uppermost leaves strongly reduced (on 
peduncles scaly), sessile, semiamplexicaul, with auriculate base. Capitula 
with 15-25(30)- florets, almost cylindrical, with fruits 17-20 mm long, in 
more or less divaricate corymbose or corymbose-paniculate inflorescence 
327 at apices of unequal peduncles, often shorter than capitula. Involucral 
bracts three- to four-rowed, sometimes with deflexed and divergent apices, 
very finely and densely pubescent dorsally, often violet; outer bracts 
oblong-ovate and cordate, inner ones linear-lanceolate or linear, subobtuse 
and with barbula of thin hairs. Corolla bright yellow, often lilac-colored 
or with lilac streaks on outer surface. Achenes ellipsoid or acute-ellipsoid, 
5-7 mm long and 2-2.5 mm wide, with one prominent longitudinal middle 
rib and two, more or less distinct, lateral ribs; mature achenes dark-brown, 
dark violet or almost black, densely covered with very short sinuate- 
transverse light-colored hairs (under high magnification!), apically 
attenuate into greenish-yellow, thin, 3 mm long beak terminating in funnel- 
shaped disk bearing pappus; outer corona of pappus of numerous short, 
lemon-colored or golden-yellow hairs (under a hand lens!), in a few or 
many rows; white hairs of inner row 7-1 1 mm long. Flowering (April) 
May to June(July). (Plate XIX, Fig. 3.) 

On outcrops of conglomerates and gypsiferous rocks, in crevices of 
granite and limestone rocks, stony and rubbly slopes. — Soviet Central 
Asiai Kyzyl-Kum, ?Amu-Darya, Pamiro-Alai, Tien Shan. General 
distribution: possibly in Afghanistan. Described from Soviet Central Asia, 
outlier mountains in Kyzyl-Kum desert. Type in Leningrad. 



324 

Note. Specimens from Tadzhikistan differ from typical specimens 
by iiaving more slender peduncles, a strongly branched corymbose 
inflorescence, slightly narrower achenes (usually dark olive-colored), and 
a few other "minor" characters. Possibly, additional materials may lead 
to splitting off the plants from Tadzhikistan as a separate race. 

2. S. crassicaulis (Trautv.) Kirp. comb. nova. — Mulgedium ? 
crassicaule Trautv. in Bull. Soc. Nat. Mosc. XXXIX, I, 2 (1866) 309 
(Enum. pi. songor No. 721). — Streptorhamphus hispidulus Rgl. in Bull.. 
Soc. Nat. Mosc. XL, II, 3 (1867) 176; O. and B. Fedtsch. Perech. Rast. 
Turk. IV, 320 (Consp. Fl. Turk. No. 2119).— Mulgedium hispidum Rgl. 
ex O. and B. Fedtsch. in Tr. Peterb. Bot. Sada XXVIII, 1 (1908) 38 (PI. 
Turkest. imprim. Alaicae... II). — Cicerbita crassicaulis (Trautv.) Beauverd 
in Bull. Soc. Bot. Geneve, 2 Ser. II (1910) 127 (seorsum impressum: 
Contrib. a I'etude des Compos. Ill, p. 29, No. 23). — C. crambifolia (Bge.). 
Beauverd f. hispidula Beauverd, op. cit. 143 (seorsum impressum: p. 45, 
No. 50). — C. chqffanjoni Beauverd, op. cit. 143 (seorsum impressum: p. 
45, No. 51). — Steptorhamphus chaffanjoni (Beauverd) O. and B. Fedtsch. 
328 in Perech. Rast. Turk. IV (1911) 319 (Consp. Fl. Turk. No. 2778).— 
Lactuca chaffanjoni (Beauverd) Danguy in Bull. Mus. His. Nat. Paris 
XX (1914) 39.— L. hispidula (Rgl.) B. Fedtsch. Rastit. Turk. (1915) 765.— 
L. chaffanjoni (Beauverd) B. Fedtsch. 1. c. 

Perennial. Root usually 1-2 cm thick, sometimes branched. Stem 
(20)40-60 cm high, sulcate, fistular, at base usually with remnants of 
petioles of previous year's leaves, glabrous or almost so in lower part, 
weakly appressed-hairy above, branched almost from base or lower third, 
producing rather numerous floriferous branches in leaf axils. Leaves 
glaucous, thick or slightly fleshy; lower leaves elliptical or oblong-ovate, 
undivided or (less often!) pinnatipartite, on rather long petioles expanded 
at base as narrow sheath, erose and with cartilaginous, more or less uncate, 
spinules along margin; middle cauline leaves oblong or spatulate, 
semiamplexicaul, cordate-auriculate at base, often more or less entire; 
upper leaves strongly reduced, usually lanceolate or linear-lanceolate, 
aristate, usually glandular-hairy; leaves on peduncles scaly, resembling 
involucral bracts, Capitula mostly with 20-25 florets, almost cylindrical 
or narrowly campanulate, with fruits 13-20 mm long and 8-12 mm wide, 
in lax corymbs at apices of slender peduncles of variable length, some 
capitula usually undeveloped. Involucral bracts often four-rowed, linear, 
linear-lanceolate, or narrowly triangular, mostly aristate, very short- and 
densely tomentose on outer surface, dorsally (mostly along midrib) covered 
with squamiform glandular hairs (also on peduncles). Corolla yellow, 
sometimes (especially after drying) lilac-colored. Achenes 6-7 nmi long 
and 2.5-3 mm wide, ellipsoid or acute-ellipsoid, with three, more or less 
prominent, ribs on each side, dark violet, covered with very short, pointed 



325 

hairs in transverse sinuate rows (under high magnification!), more distinct 
in upper part; rather sharply narrowed at apex into yellowish-green, about 
2.5 mm long beak, terminating in funnel-shaped disk bearing pappus; 
white, long hairs of pappus as long as achene or slightly longer; outer 
corona of more or less numerous, often golden-yellow, short hairs (under 
a hand lens!). Flowering May to June. (Plate XIX, Fig. 2). 

Stony and gravelly slopes at 300-2300 m. — Soviet Central Asia: 
Dzhungaria-Tarbagatai (Dzhungarian Alatau), Syr-Darya, Pamiro-Alaj, 
Tien Shan. Endemic. Described from Schrenk's collections from 
Dzhungaria (Sarybulak River). 

Note. On the label attached to one of Schrenk's authentic specimens, 
Fedtschenko, for the first time, noted that Mulgedium crassicaule Trautv. 
and Streptorhamphus hispidulus Rgl. are the same species. Thus, instead 
of accepting the more common epithet of Regel, according to the priority 
329 rule we have to select the half-forgotten name given by Trautvetter. It 
must also be mentioned that the incomplete specimens of Schrenk, which 
are not very suitable for detailed study, differ nevertheless slightly from 
typical S. hispidulus Rgl., described from the region of the extreme western 
spurs of the Dzhungarian Alatau (along the Hi River) from collections of 
P.P. Semenov. In particular, the involucral bracts in Schrenk's specimens 
are somewhat more convex and pointed and whitish on the dorsal surface 
below. Similar differences in the involucral bracts found in specimens 
collected far away from the Dzhungarian Alatau, e.g., toward the west of 
Lake Issyk-Kul, provide a basis to conclude that it is either an aging 
phenomenon or an effect of insects or lower fungi. Nevertheless, new 
complete collections are essential for absolute certainty. 

3. S. persicus (Boiss.) O. and B. Fedtsch. in Perech. Rast. Turk. IV 
(1911) 319 (Consp. Fl. Turk. No. 2776); Grossh. Fl. Kavk. IV (1934) 
258 and Opred. Rast. Kavk. 517 (sub. Streptorhampho (sic!) persico B. 
Fedtsch.) Takht. and Fed. Fl. Yerevana 339. — Lactuca persica Boiss. 
Diagn. pi. or. ser. 1, VII (1846) 9; id. in Fl. or. Ill, 806; B. Fedtsch. 
Rastit. Turk. 765. — Cicerbita persica Beauvered in BuU. Soc. Bot. Geneve, 
2 Ser. II (1910) 142 (seorsum impressum: Contrib. a I'etude des Compos. 
Ill, p. 44, No. 49); Kitam. in Res,. Kyoto Univ. Exp. Karak. a. Hinduk. 
1955, II, 405 (H. of Afghanistan, 1960). 

Perennial. Root thick, sometimes quite robust, 1-3 cm thick. Stem 
(30)40-50(60)cm high, often with remnants of petioles of previous year's 
leaves at base, sparsely puberulent (under a lens!), distinctly sulcate, 
fistular, usually branched below, thick, often assurgent with short fertile 
branches arising from leaf axils, covered with glandular hairs. Leaves 
glaucous, thick; lower leaves oblong-elliptical or lanceolate, often very 
large (up to 30 cm long, 5-7 and up to 10 cm wide), undivided or (less 



326 

often) pinnatipartite, on rather long petioles, expanded at base as narrow 
sheath, almost entire or sinuate, with cartilaginous, more or less uncinate 
spinules; middle cauline leaves few, usually sessile, semiamplexicaul, with 
auriculate or hastate base, entire or subentire, mostly oblong-elliptical; 
upper cauline leaves strongly reduced, appressed-hairy, auriculate-cordate 
at base, with branches of inflorescence or individual peduncles in their 
axil. Capitula usually with 25-35-florets, almost cylindrical or narrow 
campanulate, with fruits up to 18-25 mm long, on short, thick peduncles 
and borne in racemose-paniculate (sometimes candelabriform) 
inflorescence. Involucral bracts lax, mostly three-rowed, flat; outer bracts 
appressed hairy and glandular pubescent on dorsal surface, inner ones 
sometimes with beard of fme hairs on outer surface. Corolla yellow. 
330 Achenes acute-ellipsoid or elhpsoid, usually with one distinct rib in middle 
on each side, dark brown or yellowish, 5-7 mm long and 1.5-2.5 mm 
wide, densely covered with transversely sinuate rows of rather short hairs 
(noticeable under high magnification!), apex fairly gradually attenuate 
into dark brown beak usually longer than achene (often almost two times 
as long), terminating in somewhat funnel-shaped disk bearing pappus; 
longer hairs of pappus up to 10 mm long; outer corona of more or less 
numerous, often many-rowed, usually golden or lemon-yellow, short hairs 
(under a hand lens!). Flowering (April)May to June(July). (plate XIX, 
Fig. 4.) 

Stony and rubbly mountain slopes, on rubbly-clayey, calcareous and 
stony screes, usually at about 1000 m. — Caucasus: Southern 
Transcaucasia; Soviet Central Asia: mountainous Turkmenia (Ashkhabad 
District). General distribution: Iran Region (Iran, Afghanistan, Pakistan). 
Described from Koch's collections from southern Iran. Type in Geneva; 
isotype in Leningrad. 

Section 2. Tuberosi (Boiss.) Kirp. in Bot. Mat. Gerb. Bot. Inst. Akad. 
Nauk SSSR XXII (1963) 31S.—Lactuca sect. Tuberosae Boiss. Fl. or. Ill 
(1875) 804. — Outer corona not always distinct (under a lens!) and consists 
of almost complete row or only few isolated short hairs of same color as 
inner ones (white or transparent). A few, relatively large, transparent or 
white, erect (more or less perpendicular to achene surface), papilliform 
hairs stand out among hairs covering achene (under a lens!). Projections 
on the areola of disk usually well developed. 

Lectotype: Steptorhamphus tuberosus (Jacq.) Grossh. 

4. S. tuberosus (Jacq.) Grossh. in Fl. Kavk. IV (1934) 258 and Opred. 
Rast. Kavk. 517 (sub Streptorhampho (sic!) tuberoso (L.) Grossh.). — 
Lactuca tuberosa Jacq. Hort. bot. Vindob. I (1770) 18, t. 47; Ldb. Fl. 
Ross. II, 2, 802; Boiss. Fl. or. Ill, 806, p. p. (quoad pi. Tauricam); 
Schmalh. Fl. 11, 167, p. p. (quoad pi. Tauricam et districtum Novorossijsk); 



327 

Fedtsch. and Fler. Fl. Evrop. Ross. 1057; Stank, and Tal. Opred. Vyssh. 
Rast. (1957) 442.— '?Sonchus tuberosus Mutt. Syst. Veget. ed. 13 (1774) 
594; L. fil. Suppl. (1781) 346.— ?Lactuca sonchifolia Willd. Sp. pi. Ill, 3 
(1803) 1530, p. p. (exclusa saltern planta Cretae). — Wiestia tuberosa Sch. 
Bip. in Herb, and Winkl. Jahrb. Pharm. IV (18941) 154, in nota (vidi 
tantum seorsum impressum sub titulo "Sceleton systematis articulati 
Cichoriacearum," p. 4.). 

Perennial. Root thick, usually about 1 cm, less often up to 3 cm 
thick, mostly vertical sometimes branched. Stem 40-60(80) cm high, 
weakly and finely appressed-hairy (under a lens!), often reddish-lilac, 
slightly sulcate, usually branched in upper half. Leaves semiamplexicaul, 
densely tomentose (under a lens!); lowermost leaves early-withering (more 
331 or less undivided?); cauline leaves runcinate, petiolate or sessile, with 
irregularly arranged and unequal, cartilaginous spinules along margin; 
upper leaves strongly reduced (uppermost leaves scaly), with axillary 
peduncles, undivided, lanceolate, with large auricles at base, acuminate. 
Capitula mostly with 35^5(50) florets, oblong (younger) to cylindrical 
and campanulate, with fruits up to 30 mm long, in racemose or, less 
often (in very well-developed plants), racemose-paniculate inflorescence; 
peduncles thick, often exceeding capitula, usually arcuate above, with 
one or a few scaly leaves. Involucral brads three- or four-rowed; outer 
bracts ovate or triangular-ovate, others oblong-lanceolate or linear, often 
lilac, with distinct network of veins, more or less tomentose, along dorsal 
surface (under a hand lens!); innermost (longest) bracts attenuate at apex, 
usually with dark violet tip and lustrous stramineous inner surface. Corolla 
light blue-purple or light blue-lilac. Achenes ellipsoid, oblong, or 
lanceolate, usually with one distinct rib in center, dark violet to almost 
black, 4-5 mm long and about 2.5 mm wide, covered with transversely 
sinuate rows of very short hairs (under high magnification!), mixed with 
rather abundant, relatively long, whitish, papilliform hairs, more or less 
perpendicular to achene surface; achenes attenuate at apex, terminating 
in filiform, greenish-yellow beak, about two and one-half to three times 
as long as achene; long and white inner pappus hairs shorter than beak, 
outer corona of incomplete row of short white hairs. Flowering May to 
June(July). (Plate XIX, Fig. 5). 

Stony mountain slopes, sometimes in open montane forests. — 
European Part: Crimea; Caucasus: Western Transcaucasia (Novorcssiisk, 
Gelendzhik). The report of finding it in Georgia apparently is not correct 
and relates to the following species (5. petraeus); even more doubtful are 
the reports of it occurring outside the USSR. The typification of the species 
is unclear. The plant was described by Jacquin on the basis of specimens 
grown from seeds. The native region is not indicated in the original 
description. Possibly, the type is in Vienna. 



328 

Note. E. trautvetter (Tr. Peterb. Bot. Sada, VIII, 2, 1863, Increm. Fl. 
Phaenog. Ross. fasc. II, 520, No. 3243) reports Lactuca cretica Desf. 
from Crimea. This species is very close to S. tuberosus but differs from 
it first of all by the yellow corolla and in fact is not found in Crimea. 

5. S. petraeus (Fisch. and Mey.) Grossh. Fl. Kavk. IV (1934) 258 
and Opred. Rast. kavk. 517. — Lactuca petraea Fisch. and Mey. in DC. 
Prodr. VII, 1 (1838) 134; Ldb. Fl. Ross. II, 2, 803; Schchian in Fl. Gruz. 
VIII, 683.— Ic: Schchian, op. cit. 685, Plate 439. 
332 Perennial. Root thick, about 1 cm (less often 2-3 cm), usually vertical, 

sometimes branched. Stem (20)40-55(80) cm high, usually finely 
appressed-hairy (under a hand lens!), slightly sulcate, mostly branched 
above. Leaves semiamplexicaul, densely tomentose (under a hand lens!); 
lower leaves on long petioles, expanded below, middle leaves sessile, with 
auriculate-sagittate base, both runcinate, and with fewer cartilaginous 
spinules and usually numerous (especially in lower leaves) hairs along 
margin, also beneath on thick midrib and sometimes along other large 
veins; upper leaves strongly reduced, undivided, narrow-triangular or 
lanceolate, acuminate, basally sagittate; uppermost leaves and those on 
peduncles scaly. Capitula widi (25)35-40(50) florets, oblong (younger) 
to cylindrical and campanulate, well developed, with fruits up to (18)25- 
30(35) mm long, on somewhat thick peduncles usually longer than 
capitulum, but sometimes as long or shorter; peduncles sometimes more 
or less arcuate upward, sometimes arising obliquely, covered with short 
and dense appressed pubescence. Involucral bracts three- or four-rowed, 
outer bracts ovate or oblong-ovate, others oblong-lanceolate or linear, often 
lilac-colored, with more or less distinct network of veins, dorsally densely 
tomentose-pubescent (under a hand lens!), innermost (longest) with 
subobtuse, often apex dark violet, lustrous straw-yellow inside. Corolla 
pale yellow (according to Grossheim), on outer surface grayish-light blue. 
Achenes ellipsoid, oblong or lanceolate, dark violet to almost black, 5-6 
mm long and 2-3 mm wide, densely covered with short, transversely 
sinuate rows of hairs (under high magnification!), mixed with individual, 
relatively long, white, papilliform hairs, perpendicular to achene surface; 
achenes often truncate at base, attenuate at apex without beak, resembling 
two-toothed apex of fruits of many sedges (e.g. Carex divulsa), terminating 
in thin, light-colored beak, almost as long achene or 1.5 times as long; 
white, thin, and long inner pappus hairs almost as long as beak, outer 
corona of incomplete row of short white hairs. Flowering May to June 
(Plate XIX, Fig. 6.) 

Stony and rubbly slopes of middle mountain zone. — On screes and 
limestones, sometimes in shrub thickets. — Caucasus: Eastern and Southern 
Transcaucasia. General distribution: Armenia and Kurdistan (Turkey). 



329 

Reports about its distribution outside the USSR need confirmation. 
Described from Sovich's collections from Somkhetia. Type in Leningrad. 

Note. S. pretraeus is very close to S. tuberosus, with which it was 
often combined, for example by Boissier. Both species have much in 
common not only in their external morphology but also in the finer details 
333 of the receptacle, androecium and gynoecium. Thus, for both species the 
strong development of the antheropod^ is highly characteristic. 
Nevertheless, certain characters such as the color of the corolla, length of 
the beak in relation to the achene, presence of well-developed hairs along 
the leaf margins (which, incidentally, are also found sometimes in S. 
tuberosus) and many other "minor" characters, as well as the geographic 
range, compel me to consider S. petraeus as a separate species. 

The specimens from the Nakhichevan ASSR are shorter and differ 
by having a lilac-colored stem and involucral bracts, and also, as a rule, 
a shallow cutting of the leaves. Possibly, additional studies will provide 
evidence to treat them as a special race. In particular, the color of the 
corolla, its change with age, and the conditions of growth must be taken 
into consideration. The specimens from Kurdistan differ quite markedly 
from the type specimen by having numerous membranous hairs, which 
are slightly dilated below, on the leaf and margin of the leaf. The presence 
of a single herbarium sheet in the Herbarium of the Botanical Institute, 
Academy of Sciences of the USSR, obliges us to refrain from drawing 
any final conclusions. 

Considerable difficultly was faced in establishing the type of S. 
petraeus. In the description of Lactuca tuberosa (Prodr. VII, 1 (1838) 
133), DeCandolle cited the following locality: "... in convallibus montium 
Seidchadschi prov. Aderbeischan Persiae locis lapidorsis ... sterilibus 
(Szow. ex Fisch. and Mey. in litt.). Sovich's specimen, preserved in the 
Herbarium of the Botanical Insitute, Academy of Sciences of the USSR, 
provides an opportunity to visualize what Fischer, Meyer and DeCandolle 
understood by L. tuberosa Jacq. Judging from the dried specimen, as 
well as all other evidence, another label, in particular, "No. 183, Sonchus — 
? In lapidosis montium ad Schachbulagh' 18 maj 829, Szovits." was 
affixed by mistake to a duplicate of this same plant. In Meyer's hand are 
appended the words "Somchetiae"; ""lactuca petraea." On the same sheet, 
also evidently by mistake, the detailed original description by Meyer is 
attached. At the same time, four more sheets are preserved in the 
Herbarium of the Botanical Institute, Academy of Sciences of the USSR, 
two of which have labels written by Meyer: "In lapidosis montium versus 
Schachbulagh prov. Aberbeidschan. Szowits."; the other two sheets were 
labeled by different, still unknown persons, and have the label: "183, in 



(sic), the first letter is Russian. 



330 

lapidosis montium ad Schachbulagh, Somechetiae. 18 Mai 1829. Szovits." 
The plants under No. 183 with green divided leaves fully correspond to 
the original description of Meyer and should be considered as the type 
specimens. As regards the very first of these speciments, it cannot be 
assigned either to Lactuca tuberosa or to L. petraea. 

6. S. linczevskii Kirp. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR 
XXII (1963) 318. — Lactuca tuberosa auct. non Jacq., p. p.; Boiss. Fl. or. 
Ill, 806. — ?L. tuberosa L. f. brevirostris Krasch. in Izv. Gl. Bot. Sada 
XXIX, 1-2 (1930) 160, nom, nudum. — Steptorhamphus crambifolius auct. 
334 non Bge. p. p.; Kirp. in FI. Turkm. VII, 342.— Ic: Kirp. op. cit. 343, 
Plate LIV (sub Steptorhampho crambifolio). 

Perennial, Root about 2 cm thick, sometimes branched. Stem (25)35- 
50(70) cm high, very finely appressed-hairy (under high magnification), 
slightly sulcate, branched above. Leaves finely appressed-hairy (under 
high magnification!), especially beneath, where sometimes occasional short 
prominent hairs are still noticeable; lower leaves long-petiolate; middle 
(and upper) leaves sessile, semiamplexicaul, with auriculate or hastate 
base, both elliptical or oblong-elliptical, undivided, more or less coarsely 
toothed-spinose, less often pinnatilobate and pinnatipartite to almost 
runcinate, spinulose along margin of their lobes; uppermost cauline leaves 
narrow-triangular or triangular-lanceolate, small, those on peduncles scaly. 
Capitula with (25)30-35(40) florets, cylindrical or oblong campanulate, 
with fruits up to 30-35 mm long, in racemose or racemose-paniculate 
inflorescence; peduncles unequal, but often shorter than capitulum, thick 
(1.5-2 mm directly below capitulum). Involucral bracts somewhat 
squarrose, usually rectangular, very finely appressed-hairy dorsally (under 
high magnification!), especially in upper part, subobtuse or obtuse, often 
with violet spot and beard of very short hairs. Achenes oblong-ellipsoid 
or lanceolate, flattened, but usually with prominently convex achene, 
usually with only one distinct midrib and two less distinct lateral ribs, 
dark violet, 7-8 mm long and 2-2.5 mm wide, covered with very short, 
transversely sinuate rows of light-colored hairs (under high magnification!), 
mixed with (especially in upper part) occasional longer papilliform hairs 
more or less perpendicular to achene surface; achenes attenuate into more 
or less 1 mm long neck, bearing thin, filiform, yellowish or yellowish- 
green, 13-15 mm-long beak; longer hairs of pappus up to 12-14 mm 
long, outer crown of incomplete row or very few short light-colored hairs 
(under a hand lens!). Flowering May to July. (Plate XIX, Fig. 7.). 

Stony mountain slopes and ravines. — Soviet Central Asia: 
mountainous Turkmenia. General distribution: Iran. Described from 
Linczevsky's collections from Turkmenia (Kopetdag, vicinity of Kara- 
Kal, loldere Ravine). 



331 

7. S. czerepanovii Kirp. in Bot, Mat. Gerb. Bot. Inst. Akad. Nauk 
SSSR XXII (1963) 322. 

Perennial. Root 2-A cm thick, woody. Stem 25-45 cm high, very 
finely appressed-hairy (under high magnification!), slightly sulcate, 
branched above. Leaves finely appressed-hairy (under high magnification!), 
335 excepting uppermost, broadly or oblong-lanceolate, lamina 2-A cm wide, 
undivided (lowermost — always?), shallow-sinuate along margin and with 
rather numerous short spinules, usually acute; lower leaves on more or 
less long, semi-amplexicaul petioles, up to 15 cm long (including petiole); 
middle cauline leaves mostly sessile, with auriculate-sagittate base; 
uppermost leaves oblong or narrow-triangular, entire, with sagittate base; 
upermost leaves scaly, resembling involucral bracts. Capitula with about 
35 florets, oblong or cyUndrical, with fruits up to 25 mm long, in racemose 
or racemose-paniculate inflorescence; peduncles unequal, rather thick 
(directly below capitulum 0.5-1 mm thick). Involucral bracts somewhat 
squarrose, usually rectangular, greenish, very finely appressed-hairy 
dorsally (under high magnification!); inner bracts subobtuse or obtuse, 
with beard of short hairs at apex. Corolla color unknown. Achenes ellipsoid 
or lanceolate, with one distinct midrib and two lateral (less distinct) ribs, 
dark violet, 5-6 mm long and 2-3 mm wide, covered with very short, 
transversely sinuate rows of hairs (under high magnification!), mixed with 
large whitish papilliform hairs, borne irregularly, but more or less 
perpendicular to achene surface; achenes apically attenuate into short neck 
bearing thin, greenish, about 7 mm-long beak with disk bearing pappus; 
thin white inner hairs of pappus up to 12 mm long; outer crown like 
previous species. Flowering June. (Plate XIX, Fig. 1.) 

Stony steppe slopes in middle mountain zone. — Caucasus: Southern 
Transcaucasia (Aiotsdzor (former Daralagez) and northern part of Zangezur 
ranges; possibly, also found in Armenian SSR, in districts adjoining 
Nakhichevan ASSR). Described from collections of Tzvelev and 
Czerepanov. 

GENUS 1657. Mycelis Cass. '-^ 

Cass, in Diet. sc. nat. XXXIII (1824) 483.— Lactuca L. sect Mycelis 
DC. Prodr. VII, 1 (1838) 139.— Lactuca L. sect Ixeris Benth. in Benth. 
and Hook. f. Gen. pi. II (1873) 526, p. p. — Lactuca L. sect. Quinqueflorae 
Boiss. Fl. or. Ill (1875) 817, p.; O. Hoffm. in Pflanzenfam. IV, 5 (1894) 
372. — Cicerbita $ 3. Mycelis Beauverd in Bull. Soc. Bot. Geneve, 2 Ser. 



' Treatment by M.E. Kirpicznikov. 

^ Named by A. Cassini without explaining the etymology. 



332 

II (1910) 116 (seorsum impressum: Contrib. I'etude des Compos. Ill, p. 
18). — Lactuca subgen. Mycelis Babcock, Stebbins and Jenkins in 
Cytologia, Fujii Jubil. vol. (1937) 191, p. p. 

Capitula homogamous, few-flowered (usually comprising only three 
336 to five florets), cylindrical or narrow-campanulate, solitary on slender 
apices of branches in panicles; involucre one-rowed of a few linear or 
linear-lanceolate bracts and fewer much shorter bracts adjoining capitular 
base (considered as second row of involucre by some authors); receptacle 
usually flat, smooth; corolla yellow; achenes dark violet to almost black, 
flattened, with prominent longitudinal ribs on both sides, at apex attenuate 
into a beak terminating in broad disk; disk with very short hairs along 
margin (under a lens!); inner pappus hairs thin, fragile, slightly serrate, 
readily falling. Basic chromosome number jc = 9. 

Type of genus: Mycelis muralis (L.) Dum. 

Herbaceous plants rich in latex. According to published reports, the 
genus comprises about 30 species growing in Europe, Asia, and Africa. 

Note. As a separate genus Mycelis Cass, has not by any means been 
recognized by all investigators. Most taxonomists question the validity of 
separating Mycelis from the genus Lactuca. However, it is distinguished 
from typical Lactuca first of all by the small and usually constant number 
of florets in the capitulum as well as by the disk of the achene, which 
bears, besides the pappus, short ciliolate or papilliform hairs. It seems to 
us that the genus Mycelis, at least as expressed in its typical representative 
— M. muralis, is quite distant from the genus Lactuca and, phylogenetically 
is more allied with species of Cephalorrhynchus, from which it is 
distinguished mainly by the indistinct outer pappus, few-flowered capitula, 
and phyllaries clearly in only one series. A similar outer pappus and at 
most only a two-rowed involucre are also found in species of 
Cephalorrhynchus. Ecologically, too, M. muralis is closer to 
Cephalorrhynchus than to Lactuca subgen. Lactuca. 

1. M. muralis (L.) Dum. Fl. Belg. (1827) 60; Mikhailovskaja in Fl. 
BSSR, V, 194.— Prenanthes muralis L. Sp. PI. (1753) 797; M.B. Fl. taur.- 
cauc. II, 246, 465,. — Chondrilla muralis Lam. Fl. Fr. II (1778) 105. — 
Prenanthes parviflora Gilib. Fl. Lithuan. Ill (1781) 233; Ej. Exerc. phytol. 
I (1792) IS7.— Lactuca muralis Gaertn. De fruct. II (1791), t. CLVIII.— 
Prenanthes erysimifolia Willd. Sp. pi. Ill, (1803) 1543; M.B. Fl. taur.- 
cauc. II, 245 and III, 529.— Cicerbita muralis Walk. Sched. critic. I (1822) 
436; Beauverd in Bull. Soc. Bot. Geneve, 2 Ser. II, 139 (seorsum 
impressum: Contrib. a I'etude des Compos. III. p. 41, No. 44). — Mycelis 
angulosa Cass, in Diet. sc. nat. XXXIII (1824) 483; C.A. Mey. Verzeichn. 
Pfl. Cauc. 57.— M. muralis Rchb. Fl. Germ, excurs. II (1831) 272; Hayek 
in Hegi, 111. H. Aufl. VI, 2, 1111; Grossh. H. Kavk. IV (1934) 255 and 



333 

Opred. Rast. Kavt. 515; Klok. in Vizn. Rosl. URSR, 604; Schchian in Fl. 
Gruzii VIII, 677; Stank, and Tal. Opred. Vyssh. Rast. (1957) 444; 
337 Askerova in Fl. Azerb. VIII, 550. — Lactuca muralis Fresen. Taschenb. II 
(1833) 484; Boiss. Fl. or. Ill, 817; Popov, Ocherk Rast. i Fl. Karpat, 256; 
Kirp. in Majevski, Fl. (1954) 632. — L. muralis G. Mey. Chlor, Hannov. 
(1836) 431; Fedtsch. and Fler. Fl. Evrop. Ross. 1058; Syreistsch. 111. Fl. 
Mosk. Gub. Ill, 331. — Phoenixopus muralis W.D. Koch, Synops. Fl. Germ, 
et Helv. (1837) 430.— Lactuca muralis DC. Prodr. VII, 1 (1838) 139; 
Ldb. Fl. Ross. II, 2, 307. — L. erysimifolia DC. op. cit. 140. — L. muralis 
Less ex Schinalh. Fl. II (1897) 169.— Ic: Rchb. f. Ic. fl. Germ, et Helvet. 
XIX (1859) t. 66 (MCCCCXVII); Syreistsch. op. cit. 331; Javorka and 
Csapody, Iconogr. fl. Hungar. (1933) t. 559, fig. 4016 (sub Cicerbita 
murali); Hegi, op. cit. 1112 (photo, habitus), p. 1124 (pi. juven.), t. 227, 
fig. 4 (color.); Mikhailovskaja, op. cit. 195, Plate LXVI (ic. mala). — 
Exs..- GRF No. 973 (sub Lactuca murali (L.) Less.); PI. Polon. exs. No. 
378 (sub Mycelide murali (L.) Rchb.); Fl. Bulg. exs. (sine numero, sub 
Lactuca murali L.); Petrak, Flora Bohem. et Morav. exs. No. 1090 (sub. 
L. muralis DC); PI. Herceg. exs. No. 198 (sub. L. murali L.); Sam. PI. 
Suec. exs. ed. Hult. No. 1608 (sub L. murali (L.) Fres.). 

Perennial (?). Rhizomes not thick, often reduced, with slender, light- 
colored filiform roots. Stem mostly solitary, (15)30-100(150) cm high, 
almost wholly glabrous, smooth or, less often, somewhat sulcate, 
paniculately branched in upper part. Leaves thin, almost papery, Ught 
green above, darker or glaucous-green beneath, sometimes violet, with 
short and soft spinules along margin; lower leaves less often rosulate, on 
long-winged petioles, lyrately pinnatisect or divided, with hastate, 
triangular, rhombic or irregular lobes, of which terminal lobe larger, 
hastate, itself sometimes trilobate; middle and upper cauline leaves 
gradually reduced upward, sessile, more or less amplexicaul, with sagittate 
base, not deeply divided or almost entire. Capitula narrow-cylindrical or 
narrow-campanulate, about 10 mm long and 1.5-3(5) mm wide, solitary 
at apices of slender terminal branches of stem forming lax panicles, 
sometimes half as long as plant itself. Involucre with one distinctly 
developed row of linear or linear-lanceolate bracts; involucral bracts few 
(mostly five), subobtuse, usually with membranous border, sometimes 
reddish or violet; usually one to three smaller, more or less erect bracts 
below; involucre often deflexed after fruiting. Florets in capitulum five 
(less often four or any other number). Achenes 3-3.5 mm long (excluding 
beak) and up to 0.8 mm wide, dark violet or almost black, flattened, 
lanceolate or narrowly obovoid, with more or less numerous prominent 
longitudinal ribs, attenuate above into neck, terminating in greenish-yellow, 
short, up to 1 mm long beak with broad disk bearing pappus; disk densely 
covered with very fine white hairs (under high magnification!); 



334 

338 achenes covered with numerous pointed, transparent, upward-directed 
(under a lens!); inner pappus hairs fragile, very thin, somewhat barbed, 
readily detached, white; outer hairs of pappus short (under high 
magnification!), not always distinct. 2n = 18. Flowering June to August 
(September). (Plate XX, Fig. 9.) 

Shady and moist coniferous, deciduous and mixed forests, forested 
ravines, on humus-rich nonacidic soils; on wet rocks and old delapidated 
stone walls; in mountains up to upper limit of forests; in the Caucasus 
mistly in upper forest and subalpine zones at 1,300-2,300 m. European 
Part: Baltic region, Ladoga-Ilmen, Upper Volga, Volga-Kama (south), 
Upper Dnieper, Middle Dnieper, Volga-Don,Upper Dniester, Bessarabia, 
Crimea, Lower Don; Caucasus: all regions. General distribution: 
Scandinavia, central Europe, Atlantic Europe, Mediterranean, Balkans- 
Asia Minor, North America (introduced). Described from Europe. Type 
in London. 

Note. Some authors consider Prenanthes vulgaris Gueldenst. as a 
synonym of M muralis. According to the citations of Ledebour and "Index 
Kewensis," this name was published by Gueldenstedt in his "Reisen durch 
Russland und in Caucasischen Gebirge" (Vol. I (1787) 113). After 
carefully examining this paper, I did not find there any other name besides 
P. muralis. 

The question of the duration of M. muralis is not clear and needs 
confirmation. Many authors, despite the presence of a rhizome, consider 
it an annual. 

Economic Importance. There are reports that the leaves of M. muralis 
are suitable for making salads. 

GENUS 1658. Cephalorrhynchus Boiss. ' ^ 

Boiss. Diagn. pi. or. ser.l, IV (1844) 28. — Lactuca sect. 4. Cicerbita 
Benth. in Benth. and Hook. f. Gen. pi. II (1873) 526, p. min. p. — Cicerbita 
§ 4. Cephalorrhynchus Beauverd in Bull. Soc. Bot. Geneve, 2 ser. II 
(1910) 117 (seorsum impressum: Contrib. a I'etude des Compos. HI, p. 19). 

Capitula homogamous, 10-15(25)-flowered, small (with fruits up to 
(7)10-18 nun long), mostly cylindrical or oblong, usually in corymbose 
or corymbose-paniculate, inflorescence, less often solitary at apices of 
long slender branches. Involucre (excluding few reduced leaves on 
peduncle, often very close to it), three-rowed, less often two- or four- 
rowed, involucral bracts herbaceous, densely covered dorsally with short 
papillae (under a hand lens!); innermost (and longest) bracts with beard 



' Treatment by M.E. Kirpicznikov. 

^ From the Greek words cephale — head, and rhynchos — beak, nose. 



335 

of thin hairs at apex (under a hand lens!). Receptacle smooth, flat. Anthers 
oblong-linear with appressed sagittate basal appendage. Style and its 
branches densely covered with upward-directed, light-colored hairs. 
Achenes usually narrow or Unear-fusiform, often slightly curved, with 
339 more or less numerous prominent longitudinal ribs, usually with pointed, 
upward directed, short hairs (under a hand lens!); besides, very short 
hairs (under high magnification); in transversely sinuate rows; beak thin, 
of different color than rest of achene. Pappus of outer row forming corona 
of short (hardly visible without lens) hairs, firmly attached to disk and 
inner row of relatively long, usually as long as achene, rather thin, white, 
fragile, usually very readily detached hairs, appearing smooth (under high 
magnification, barbed). 

Type of genus: Cephalorrhynchus glandulosus Boiss. 

The genus comprises about 10 species of biennial or perennial herbs 
with thickened or tuberous underground parts. Plants mainly growing in 
Transcaucasia and Soviet Central Asia, occasionally in Crimea; outside 
the USSR, known definitely from Iran, Turkey, Syria, Lebanon, and 
Afghanistan. 

Boissier, who established the genus Cephalorrhynchus considered 
it quite close to the genus Lactuca, differing from the latter mainly by 
having cylindrical or fusiform, not compressed, achenes. Bentham, 
Hoffmann, and many others included Cephalorrhynchus in Lactuca, while 
Bower united it, as a section, with Cicerbita. Actually, the genus 
Cephalorrhynchus may be closest to the genus Scariola. 

1. Upper part of stem and especially peduncle densely covered with 
glandular hairs 2. 

+ Stem in upper part and also peduncles smooth or almost smooth, 
only sometimes with small number of hairs 4. 

2. Stem wholly (sometimes excluding only lowest part) covered with 
squamiform glandular hairs. Plants found mainly in the Caucasus, 
sometimes in Crimea 3. 

+ Stem glandular-hairy only in upper half, smooth or almost smooth 

below. Plant endemic to Tien Shan 

6. S. subplumosus S. Kovalevsk. 

3. Lower cauline leaves pinnatisect, with large number of almost equal 
lateral segments. Plant found in the Nakhichevan ASSR 

3. C. kirpicznikovii Grossh. 

+ Lower cauline leaves pinnatipartite or pinnatisect to lyrately 
pinnatisect, with small number of lateral segments and distinctly 
larger terminal segment. Plant found in the Caucasus and Crimea 

1. C. tuberosus (Stev.) Schchian 

4(1) Stem leafless or almost so at fruiting; capitula solitary at apices of 



336 

long slender branches (peduncles) 

8. C. polycladus (Boiss.) Kirp. 

340 + Plants leafy at fruiting; capitula in paniculate inflorescence 5. 

5. Leaves (except uppermost) undivided, broadly elliptical or oblong- 
obvate, sessile, glaucous, somewhat fleshy, with prominent veins 
beneath; beak of achenes up to 1 mm long. Plants endemic to 
southern Transcaucasia (Daralagez, Nakhichevan ASSR) 

7. C. takhtadzhianii (Sosn.) Kirp. 

+ Leaves of different shape; beak of achenes usually longer 6. 

6. Lower leaves long (including petiole often exceeding 20 cm in 
length), wide (in broadest parts up to 8-15 cm), pinnatisect, with 
large trilobate terminal segment. Plants endemic to Talysh 

2. C. talyschensis Kirp. 

+ Lower leaves shorter, or weakly divided to undivided 7. 

7. Beak half as long as achene, sometimes slightly shorter, bifid in 
lower part as if apical; pappus of very thin hairs, often persistent 
on mature achenes 4. C. soongoricus (Rgl.). S. Kovalevsk. 

+ Beak up to one-third as long as achene, not bifid; pappus of very 

thin, extremely fragile hairs. Plants endemic to Turkmenia 

5. C. kossinskyi (Krasch.) Kirp. 

Section 1. Cephalorrhynchus. — Stem leafy; capitula in corymbose- 
paniculate inflorescence; achenes mostly dark violet with thin, usually 
greenish-yellow beak. 

Type of section: type of genus. 

1. C. tuberosus (Stev.) Schchian in Zam. po Sist. i Geogr. Rast. 
Tbil. 23 (1963) 99. — Prenanthes tuberosa Stev. in Mem. Soc. Nat. Mosc. 
IV (1813) 99; M.B. Fl. taur.-cauc. Ill, 529.— P. hispida M.B. Fl. taur.- 
cauc. II (1808) 245, non Pall. 1771 (quae revera planta ex affinitate 
Chondrilla est). — Cephalorrhynchus hispidus Boiss. Fl. or. Ill (1875) 821; 
Grossh. Fl. Kavk. IV (1934) 259 and Opred. Rast. Kavk. 517; Schchian 
in Fl. Gruzii, VIII, 692; Askerova in Fl. Azerb. VIII, 560.— C. confertus 
Conrath and Freyn ex Freyn in Bull. Herb. Boiss. Ill, 9 (1895) 477; 
Grossh. Fl. Kavk. IV (1934) 259 and Opred. Rast. Kavk. 5\1 .—Cicerbita 
conferta Beauverd in Bull. Soc. Bot. Geneve, 2 Ser. II (1910) 140 
(seorsum impressum: Contrib. a I'etude des Compos. Ill, p. 42, No. 46). — 
C. hispida Beauverd, op. cit. p. 141 (seorsum impressum: p. 43, No. 
48). — Steptorhamphus hispidus Bomm. in Beih. z. bot. Centralbl. LX, 
Abt. B (1939) 224.— Lactuca hispida auct. non DC.: Ldb. Fl. Ross. II, 2, 
807; Schmalh. Fl. II, 169; Fedtsch. and Fler, Fl. Evrop. Ross. 1059. 

341 Biennial or perennial. Root tuberous. Stem (25)50-75(105) cm high, 
fistular, finely sulcate, branched above, densely covered with squamiform- 



337 

glandular hairs. Leaves thin, more or less glaucous beneath; lower leaves 
10-15(25)cm long, with petioles (2-3 mm wide) expanded at base and 
semiamplexicaul, lamina pinnatipartite, pinnatisect or lyrate, with large 
triangular terminal lobe, coarsely sinuate-toothed, with spinescent teeth 
(lower leaves often lacking in fertile specimens); leaves gradually reduced 
toward stem apex, with shorther petioles, expanded at base and auriculate- 
cordate or sagittate, more or less lacking teeth, acuminate; upper cauline 
leaves sessile, strongly auriculate-cordate and sagittate base, broadly 
lanceolate to oblong, oblong-lanceolate and narrow-triangular; uppermost 
leaves linear or narrowly triangular-linear, scaly at base of peduncles and 
especially on peduncles. Capitula usually wirh 12 florets, cylindrical or 
oblong-elongated, with fruits about 1 5 mm long, in corymbose-paniculate 
inflorescence; peduncles rather slender, usually shorter than capitulum, 
more or less densely covered with squamiform-glandular hairs. Involucral 
bracts usually two-rowed, narrow-triangular or linear with membranous 
border, dorsally (under a hand lens!) densely papillose, with slightly 
carinate projection in middle; inner bracts obtuse, usually with dark violet 
spot and barbula of very short and thin hairs at apex. Corolla yellow, 
usually blue after drying. Achenes narrow^ or linear-ftisiform, with 5-15 
longitudinal ribs (more distinct in upper part), dark violet, about 5 mm 
long and 0.5-1 mm wide, covered with very short, transversely sinuate 
rows of hairs (under high magnification!); and upward-directed hairs in 
upper part (under a hand lens!); attenuated upper part of achene 
terminating in thin, greenish-yellow, 2-3 mm long beak (very rarely dark 
brown); outer row of pappus (crown) dense, of short hairs, inner one of 
rather thin, white, fragile, readily detached, 5-6 mm long hairs. Flowering 
May to July. (Plate XX, Fig. 2.). 

Forests and forest edges, in thickets, on shady slopes. — European 
Part: Crimea; Caucasus: Eastern, Western and Southern Transcaucasia. 
General distribution: Armenia and Kurdistan, Iran Regim (Iran). Described 
from Georgia on the basis of collections of H. Steven. Type in Leningrad. 

Note: The authors of Index Kewensis and Boissier placed an equals 
sign between Cephalorrhynchus hispidus (M.B.) Boiss. [i.e., C. tuberosus 
(Stev.) Schchian} and Lactuca microcephala DC. (Prodr. VII, 1, 1838, 
p. 134), the species described by DeCandolle from Iran from Aucher's 
collections (Fl. exs. No. 3517). In the opinion of A. Schchian, however, 
these are different species, and, accordingly, she made the new combination 
342 Cephalorrhynchus microcephalus (DC.) Schchian {Fl. Gruzii VIII (1952) 
693). She also pointed out the identity of C. confertus with the earlier 
described C. hispidus (i.e., C. tuberosus). 

C. tuberosus is a markedly variable species in the growth vigor of the 
overall inflorescence (corymbose panicles), number of longitudinal ribs on 
the achene, and also the degree of pubescence on the stem and peduncles. 



338 

2. C. talyschensis Kirp. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk 
SSSR XXII (1963) 311.— Lactuca deltoidea C.A. Mey. Verzeichn. Pfl. 
Cauc. (1831) 56, p. p. (solum quod plantas talyschenses attingit); Ldb. 
Fl. Ross II, 2, 806, p. p.; Boiss. Fl. or. Ill, 816, p. p. — Cicerbita deltoidea 
auct. mult. fl. Cauc. p. p. (plantae e Talysch): Grossh. Fl. Kavk. IV (1934) 
252 and Opred. Rast. Kavk. 514; Schchian in Fl. Gruzii, VIII, 667; 
Askerova in Fl. Azerb. VIII, 545. 

Perennial. Rhizome?. Stem (60)70-80(100) cm high, solitary, fistular, 
slightly sulcate, smooth or almost so, paniculately branched above. Leaves 
large, thin, green above, glaucous beneath, mostly covered (especially 
along veins) with pale, very narrow squamiform hairs; lower leaves 
pinnatisect, with large, more or less trilobate, apical segment, with opposite 
or alternate, oblong, ovate-triangular or irregular lateral segments, petiole 
more or less winged, semiamplexicaul, expanded at base; upper leaves 
smaller, with weakly cut up laminas, at base often auriculate; uppermost 
leaves small, oblong-lanceolate to linear, scaly on peduncles. Capitula 
broadly cylindrical to campanulate, (10)13-15 mm high on peduncles, 
usually exceeding capitula; inflorescence paniculate, sometimes large (30- 
40 cm long, 10-15 cm wide in broadest part). Involucre, usually three- 
rowed; inner involucral bracts oblong-linear, two to three times as long 
as outer, at tip usually violet, roundish, with barbula of very short and 
thin, light-colored hairs (under a hand lens!). Florets in capitulum about 
15; corolla yellow (?). Achenes 6-6.5 mm long and 1 mm wide, linear- 
fusiform, with 7-10 prominent ribs on each side, mature achenes dark 
violet or reddish-brown, covered with short, stiff, upward-directed, light- 
colored hairs (under a hand lens!), in somewhat sinuate rows; achenes at 
apex attenuate into yellowish, distinct, 1-1.5 mm long" beak; outer row of 
pappus hairs dirty-white forming crown; inner hairs snow-white, very thin, 
6-7 mm long readily detached. Flowering June. (Plate XX, Fig. 3.) 

Middle mountain zone, in forests. — Caucasus: Talysh. Endemic. 
Described from Meyer's collections from Talysh. Type and isotype in 
Leningrad. 

Note. Freyn identified the plant collected by Sintenis (Iter orientale 
1894, No. 7210; Armenia turcica, Szandschak Giimiischkhane, Tempede, 
343 in Saxosis,19, VII) in Turkey as Lactuca deltoidea C.A. Mey. (i.e., my 
species Cicerbita talyschensis Kirp.). Beauverd (op. cit. p. 129) agreed 
with Freyn's identification. Actually, this plant, which I had the opportunity 
to study (only one specimen), although it resembles C. talyschensis, is 
distinguished at the same time by the larger achenes, leaves, and also 
involucral bracts. Apparentiy, a separate race of the C talyschensis alliance 
grows in Turkey. 

3. C. kirpicznikovii Grossh. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk 



339 

SSSR XIII (1950) 31; Opred. Rast. Kavk. 517; Askerova in Fl. Azerb. 
VIII, 560 ("C kirpitschnikowii"). 

Biennial or perennial. Root tuberous, mostly globose, 15-20 mm thick. 
Stem (15)30-55(80) cm high, fistular, often irregularly sulcate, branched 
above, usually glabrous near base, with increasingly dense pubescence of 
squamiform-glandular hairs above. Leaves thin; lower leaves up to 25 
cm long, long-petiolate, petiole usually longer than lamina, at base 
expanded, with broadly elliptical or broadly obovate lamina, pinnatisect 
into oblong or oblong-linear, lateral segments, which are almost equal 
and similar in shape and more or less horizontal, entire, shallow-toothed, 
with small number of subobtuse spinules along margin and apex; middle 
cauline leaves smaller than lower ones, with petiole strongly expanded at 
base, and auriculate-sagittate, almost amplexicaul, toothed along margin; 
upper cauline leaves sessile or subsessile, relatively small, with lamina 
often only with three to five segments (or lobes), antrorse and acute, or 
lamina more or less entire; uppermost leaves narrowly triangular, strongly 
reduced, scaly on peduncles. Capitula mostly wirh 15-florets, cylindrical 
or oblong, 13-17 mm high, in corymbose or (less often) corymbose- 
paniculate inflorescence; peduncles slender like inflorescence branches, 
rather densely covered with squamiform glandular hairs. Involucral bracts 
usually two-rowed, with membranous border, dorsally densely covered 
with short papillae (under high magnification), usually with carinate 
projection in middle (as if along midrib), with few squamiform-glandular 
hairs, inner bracts usually with violet spot and beard of very thin and 
short light-colored hairs (under a lens!). Corolla pale yellow. Achenes 
narrow- or linear-fusiform, usually with numerous (about 15) longitudinal 
ribs, brown-dark brown to dark violet and blackish, about 6 mm long 
and up to 0.5 mm wide, covered with short hairs in transverse sinuate rows 
(under a hand lens!); achenes gradually attenuate at apex bearing light- 
colored, yellowish, about 3 mm-long beak; pappus almost indistinguishable 
from former species, but with 6-7 mm long hairs of inner row, not so 
easily detached. Flowering May to June. (Plate XX, Fig. 1.). 
344 On stony slopes, in shade of rocks. Shrub thickets and sometimes in 
open montane forests at 900-2600 m. — Caucasus: Southern Transcaucasia 
(Nakhichevan ASSR). General distribution: Probably growing in northern 
Iran. Described from Nakhichevan ASSR. Type and isotype in Leningrad. 
Note. We have tentatively referred two incomplete specimens from 
the Armenian SSR to C. kirpicznikovii: one of them was collected in the 
Megri District and does not have fertile parts; the other from the 
Echmiadzin District, has only immature capitula. Both are distinguished 
from typical C. kirpicznikovii by certain details of the form of the leaf 
blade, but mainly by the presence of a fairly large number of membranous 
glandular hairs on the lower surface of the blade (especially on the upper 



340 




345 Plate XXI. 

1 — Cicerbita thianschanica (Rgl. and Schmalh.) Beauverd, habit of plant, achene; 

2 — C. alpina (L.) Wallr., achene; 3 — C. azurea (Ldb.) Beauverd, achene; 

4 — C. uralensis Beauverd, achene; 5 — C. macrophylla (Willd.) Wallr., achene; 

6 — C. sevanensis Kirp., achene; 7 — C. racemosa (Willd.) Beauverd, achene; 

8 — C. prenanthoides (M.B.) Beauverd, achene; 9 — C. bourgaei (Boiss.) Beauverd, achene. 



341 

cauline leaves), which are more abundant along midrib and the more 
obvious lateral veins; these hairs are present in much larger number on 
the involucral bracts of the Echmiadzin specimen than is characteristic 
for C. kirpicznikovii. It is quite possible that a special race ofthe genus 
Cephalorrhynchus of the C. kirpicznikovii alliance grows in the Armenian 
SSR, but this question can be finally resolved only by studying more 
complete material. 

Grossheim compared C. kirpicznikovii with C. hispidus (M.B.) Boiss. 
[i.e., C. tuberosus (Stev.) Schchian]. In fact, it is better to compare this 
species with C. candolleanus Boiss., to which it actually is very similar. 
According to Kovalevskaja (compare her labels on the herbarium sheets), 
C. kirpicznikovii should generally be identified with C. candolleanus. Such 
an inference is hardly justified. Beauverd (1. c.) already wrote that the 
peduncles of C. candolleanus are weakly glandular-hairy or entirely 
eglandular, whereas the peduncles of C. kirpicznikovii are characterized 
by abundant glandular hairs. Moreover, C candolleanus, according to 
the analysis of Beauverd, is distinguished by the relatively smaller size 
of the achenes and the inner row of the pappus. However, it must be 
noted that the material identified as C candolleanus is not uniform. 
Apparently, Boissier combined at least two quite distinct species under 
this name. One of them is represented by the specimens of Heldreich 
from Lycia (In lapidosis sylvaticis montis Solyma, Lycia, Heldreich, Majo 
1845); these are shorter plants with almost rhombic lateral segments of 
the lower leaves, which often have a relatively large triangular apical 
segment. The specimens from Syria ought to be assigned to another 
346 species; only two poor specimens collected by Boissier (Syria, Antilibanus 
ad Cedros. E. Boissier, Junio 1846) could be examined from this region. 
Superficially, they strongly resemble C. kirpicznikovii, but they can be 
distinguished easily at least by the little pubescence of the peduncles and 
still more weakly pubescent stem. Probably, the name C. candolleanus 
Boiss. should be retained for the Syrian specimens, and the Heldreich' s 
specimens should be considered the type specimens of a separate species. 
The specimen of Sintenis and Rigo {Iter cyprium 1880, No. 798, 18 
VI) is of great interest. In habit, it is very similar to the depauperate 
specimens of C. kirpicznikovii but can be easily distinguished by much 
weaker pubescence on the peduncles as well as by the light brown color 
of the achenes with a concolorus beak. Finally, the specimens of 
Bommiiller {Iter Syriacum II (1910) No. 12061) from northern Lebanon, 
identified by him as Cicerbita candolleana (Boiss.) Beauverd, are 
characterized by long and slender, often smooth peduncles. All this raises 
doubt about the integrity of Cephalorrhynchus candolleanus as a distinct 
species. However, a definitive conclusion can be drawn only from more 
extensive material than what is at my disposal now. 



342 

4. C. soongoricus (Rgl.) S. Kovalevsk. in Fl. Uzb. VI (1962) 491. 
— Lactuca soongorica Rgl. in Bull. Soc. Nat. Mosc. XL, II, 3 (1867) 
174; O. and B. Fedtsch. Perech. Rast. Turk. IV, 320 (Consp. Fl. Turk. 
No. 2782); Popov in Tr. Uzb. Gos. Univ. Nov. Ser. No. 27, Biol. Vyp. 
14, 104. — Cicerbita soongorica (Rgl.) Krasch in Nevski in Tr. Bot. Inst. 
Akad. Nauk SSSR, Ser. 1, 4 (1937) 294 ["C. soongarica (Rgl.) H. 
Krasch."]; Kirp. in Fl. Turkm. VII, 328.— Ic: S. Kovalevsk., op. cit. t. 
XLVIII, Fig. 1. 

Perennial. Plants with thick tuberous base. Stem solitary, 50-80 cm 
high, sulcate, sometimes slightly violet at base, glabrous or almost so, or 
with more or less large number of squamiform hairs below. Leaves tender, 
green, oblong-obvate or lanceolate, undivided and more or less strongly 
sinuate to lyrate and pinnatisect, usually spinose-toothed; lower cauhne 
leaves long-petiolate, middle cauline leaves sessile, amplexicaul, with 
auriculate or sagittate base; uppermost leaves strongly reduced, scaly in 
inflorescence. Capitula, usually with 10-15-florets, including fruits 9-12 
mm long, cyhndrical, sometimes slightly broadened upward, mostly on 
long and slender peduncles, in compound paniculate inflorescence. 
Involucral bracts usually three-rowed, narrow, more or less acute, often 
violet or with violet spot, very finely pubescent on outer side (under a 
hand lens!), narrowly membranous along margin; innermost bracts often 
with apical barbula of hyaline hairs. Corolla light blue. Achenes narrow- 
fusiform, dark brown-red or dark violet to almost black, about 5 nrni 
long and 0.5 wide, usually with five prominent longitudinal ribs on each 
side, covered with inconspicuous, very short, light colored, upward-directed 
hairs (under a hand lens!); beak thin, greenish, up to half as long as 
347 achene, sometimes only slightly shorter than achene, slightly bifid below 
as if mounted on achene; inner pappus hairs often persistent on mature 
achene, almost as long as achene. Flowering April to June(July). (Plate 
XX, Fig. 4) 

In scrub-forest zone. — pistachio and juniper forests, in more or less 
open places — on stony and clayey slopes in mountain ravines, on 
gypsiferous deposits; up to 2,000 m; found in the Kirgiz Alatau (according 
to Nikitina, 1960) in dense coniferous forests with thick moss cover. — 
Soviet Central Asia: Balkhash (southern part, rather less frequent, 
Dzhungaria-Tarbagatai (small collections from Dzhungarian Alatau, Chulak 
Mountains), Pamiro-Alai, Tien Shan. Endemic (?). (Probably found in 
Afghanistan.) Described from Ludwig's collections ("Kirghisensteppe") 
and Semenov's from Trans-Ili Alatau. Lectotype and paratype in 
Leningrad. 

Note. The specimen with the following label in Regel's hand, 
''Lactuca soongorica Rgl. teste Rgl. leg. Semenow am Flusse Assy in 
Alatau transiliensis 5000," has been selected as the lectotype. 



343 

5. C. kossinskyi (Krasch.) Kirp. comb. nova. — Cicerbita kossinskyi 
Krasch. in Izv. Gl. Bot. Sada XXVI, 2 (1927) 115; Kirp. in Fl. Turkm. 
VII, 328. — Lactuca kossinskyi Krasch. i.e. in synon. — Ic: Kirp. op. cit. 
331, t. LII. 

Perennial. Root thick, fusiform, mostly branched. Stem 30-70(100) 
cm high, at base 5-10 mm thick, sulcate with sparse, simple, narrow- 
squamiform or glandular hairs (under a hand lens!), with loose and spongy 
tissue inside. Leaves somewhat fleshy, with thick midrib and more or 
less prominent network of lateral veins, erose-toothed along margin and 
with individual cartilaginous spinules; basal leaves large, 10-20 cm long 
and 5-7 cm wide, oblonge-elliptical or oblong-obovate, narrowed into 
broad and long-winged petiole; middle cauline leaves gradually reduced, 
obovate or lanceolate, usually sessile, semiamplexicaul; uppermost leaves 
sessile, with auriculate base, usually entire; leaves on peduncles small to 
scaly, at least on outer side gladular and tomentose. Capitula with 15-25- 
florets, with fruits 15-18 mm long, oblong or broadly lanceolate, mostly 
on long peduncles, borne in lax corymbose-paniculate inflorescence. 
Involucral bracts three- or four-rowed, narrow, with more or less dense, 
but very short, glandular pubescence on outer surface; outer bracts 
triangular or lanceolate, often with distinct midrib; innermost bracts linear- 
lanceolate to linear, at apex with beard of hyaline tomentose-entangled 
hairs. Corolla blue. Achenes dark brown or reddish-brown, about 6 mm 
long and 0.5 mm wide, fusiform, with prominent longitudinal ribs covered 
348 with very short, stiff, upward-directed hairs; achene apex conically 
narrowed forming a light-colored, mostly yellowish or greenish-yellow 
beak, about one-third as long as achene; inner hairs of pappus almost as 
long as achene, readily detached. Flowering May to July. (Plate XX, Fig. 5) 

In ravines, among stones, in more or less shady places. — Soviet 
Central Asia:, mountainous Turkmenia. Endemic. Described from 
Turkmenia. 

Note. From time to time specimens are found in which the lower 
leaves are runcinate — var. runcinata (Krasch.) Kirp. 

Krascheninnikov did not select a type for the species. The specimen 
with the following label is, therefore, proposed as the lectotype: 
Transcaspian Region, Ashkhabad District, Kopetdag Mountains, Firyuza. 
V. Lipsky, 3.V.1912, No. 1516 (V.I. Lipsky, Ekspeditsiya v Zakaspiiskuyu 
Oblasf 1912). [Expedition to the Transcaspian Region, 1912]. Lectotype 
and other type materials preserved in Leningrad. 

6. C. subplumosus S. Kovalevsk. in Bot. Mat. Gerb. Inst. Bot. Acad. 
Nauk Uzb. SSR XV (1959) 53; eadem in Fl. Uzb. VI, 491. 

Perennial. Roots tuberous. Stem 5-10 cm high, fistular, finely sulcate, 
branched above, finely appressed-hairy below (under high magnification!) 



344 

or almost glabrous, mixed with more or less numerous, squamiform, 
glandular hairs above. Leaves thin; basal and lower cauline leaves, early- 
withering; middle cauline leaves on broad, winged, basally auriculate, 
semiamplexicaul petiole, usually with lyrately pinnatipartite or pinnatisect 
lamina, with large terminal segment; upper cauline leaves like middle, 
but usually with one pair of lobes, short-petiolate or sessile; leaves abruptly 
narrowed at base of fertile branches from broad, auriculately amplexicaul 
base; leaves on peduncles reduced, resembling involucral bracts. Capitula 
usually with 10-florets, cylindrical or oblong, with fruits (well developed!) 
about 10 mm long, in corymbose-paniculate inflorescence; peduncles 
slender, like inflorescence branches, densely covered with squamiform- 
glandular hairs. Involucral bracts mostly two-rowed (excluding few leaflets 
on peduncles, closely adjoining involucre), narrow-triangular or linear, 
dorsally densely covered with short papillae (under high magnification!); 
inner bracts longer, usually with carinate projection in middle, mostly 
with more or less numerous, squamiform, glandular hairs, apically with 
beard of short, light-colored hairs (under a hand lens!). Corolla white (?). 
Achenes narrow-fusiform, usually slightly curved, with numerous (about 
15) longitudinal ribs, light brown, dark brown or brown-dark brown, 4- 
5 mm long, covered with transverse sinuate rows of short, fme, upward- 
directed hairs, more distinct in upper part (under a lens!); apically achenes 
349 attenuate into about 1.5 mm long, yellowish-green beak; inner row of 
pappus hairs fragile, easily detached, barbed (slightly plumose); (under 
high magnification !), almost as long as achene. Flowering June to July. 
(Plate XX, Fig. 6.) 

Middle mountain zone, near streams, under trees and in juniper forests. 
— Soviet Central Asia: Tien Shah (western part). Endemic. Type and 
isotype in Tashkent. 

Note. The specimen labled by Pavlov as "Mulgedium alboligulatum 
N. Pavl. sp. n. Bostandyk, Ugamsk Range: Boguchal-sai Ravine, 1,600 
m, gravelly slopes. 26.VI.1954. No. 62, Leg. ? V.N. Pavlov" should also 
be referred to this species. 

7. C. takhtadzhianii (Sosn.) Kirp. Comb. nova. — Lactuca 
takhtadzhianii Sosn. in Soobshch. Akad. Nauk Gruz. SSR II; 7 (1941) 
653; Grossh. Opred. Rast. Kavk. 516; Askerova in Fl. Azerb. VIII, 553. 
— Ic: Sosn op. cit. p. 655. 

Biennial or perennial. Rhizome lignified, sometimes branched. Stems 
solitary or two to several, 13(20-40)50 cm high, distinctly sulcate, usually 
appressed-hairy or flocculose-pubescent below, with more or less 
considerable mixture of short squamiform hairs (under a hand lens!) 
branched almost from middle. Leaves, except uppermost, broadly or 
oblong-obovate, or broadly elliptical, glaucous, somewhat fleshy, unequally 



345 

spinose-toothed, with straight or more or less strongly bent teeth, with 
prominent veins beneath, sessile, semiamplexicaul and auriculate-sagittate 
at base; leaves near base of large inflorescence branches reduced, more 
or less ovate or lanceolate, on peduncles scaly. Capitula with about 15- 
florets, cylindrical or turbinate-cylindrical, about 10 mm high, in 
corymbose-paniculate inflorescence; peduncles slender, usually sparsely 
sericeous. Involucre three- or four-rowed; involucral bracts, at least partly, 
reddish, dorsally covered with dense, short, papilliform hairs (under a 
hand lens!); inner bracts with membranous border, reddish at apex, 
subobtuse or obtuse, with beard of fine, silky, slightly entangled white 
hairs. Corolla dirty blue. Achene about 4.5 mm long and up to 1 mm 
wide, narrowly elliptical or narrowly oblong-obovoid, dark reddish-brown, 
more or less flattened, usually slightly curved, mostly with five prominent 
longitudinal ribs on each side, with transversely sinuate rows of numerous, 
very short, light-colored hairs (under a hand lens!); apically achenes 
slightly attenuate, terminating in small, slightly roundish notch, bearing 
distinct, thin, but short (0.5-1 mm long), greenish beak; inner pappus 
hairs fine, white, silky, about 4 mm long, more or less firmly attached to 
disk. Flowering June to August. (Plate XX, Fig. 7.) 
550 Dry, clayey, rubbly and stony slopes in the middle mountain zone. — 

Caucasus: Southern Transcaucasia (Daralgyoz, Nakhichevan ASSR). 
Endemic. Described by Sosnowsky from Takhtadjan's collections. Type 
in Yerevan. 

Note. Apparently, a quite rare plant. There is good material in the 
Herbarium of the Botanical Institute, Academy of Science of the USSR, 
which was identified by An. A Fedorov (collections of 1955, in the 
Shakhbuz District of the Nakhichevan ASSR). We have prepared the 
description from this material, which, without doubt, belongs to C. 
takhtadzhianii. It does not fully correspond to the original description, 
which, in all probability, is due to the fact that D.I. Sosnowsky had at his 
disposal poor or immature specimens. 

The underground parts, more or less rounded or slightly fleshy leaves, 
very short achene beak, and corolla color are all characters that clearly 
differentiate C. takhtadzhianii from the other species of Cephalorrhynchus. 
In habit, this plant more or less resembles Lactuca rosularis. Specimens 
with carefully unearthed underground parts and an ample number of 
mature achenes are extremly essential for confirming the taxonomic status 
of this species. 

Section 2. Zollikoferiastrum Kirp. sect, nova in Addenda XXVIII, 
725. — Stems branched from base, more or less leafless at fruiting; capitula 
solitary at apices of long, slender branches (peduncles); achenes light 
colored, terminating in short dark beak. 



346 

Type of section: Cephalorrhynchus polycladus (Boiss.) Kirp. 

8. C. polycladus (Boiss.) Kirp. comb. nova. — Zollikoferia polyclada 
Boiss. Fl, or. Ill (1875) 827. — Launaya polyclada Kuntze, Rev. Gen. I 
(1891) ?)5\.—Launaea polyclada Burk. Fl. PI. Baluch. (1909) 44 (non 
vidi). — Cicerbita lipskyi Krasch. in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 

I, 3 (1936) 353. — Lactuca lipskyi Krasch. I.e. nom. in synon. — L. 
intricatissimi Rech. f. in Dan. Biol. Skr. 8, 2 (1955) 207 (Symbolae 
Afghanicae II); Kitam. in Res. Kyoto Univ. Exp. Karak. a Hinduk. 1955, 

II, 431 (Fl. of Afghanistan, 1960). — Launaea polyclada B. Fedtsch. in 
herb, and ex Kirp. in Fl. Turkm. VII (1960) 326.— Ic. Rchb. f. op. cit. 
207, fig 143 (sub Lactuca intricatissima: habitus) and 209, fig. 145 (infra 
a sinistra — calathidium, flos, achenium). 

Perennial. Semishrub, (15)25-45 cm high, branched from base, with 
numerous glaucous, weakly sulcate or striate branches becoming slender 
toward apex. Leaves slightly thickened, glacuous, finely appressed-hairy 
(under a hand lens!), crowded mainly in lower part of stem, almost entirely 
withering before fruiting; basal leaves 2-6 cm long, lyrate, with relatively 
large oblong or roundish-ovate terminal segment and somewhat small 
351 lateral segnients, sinuate-toothed with narrow-winged, long, almost 
amplexicaul petiole; cauline leaves (often absent!) lyrate or linear, 
semiamplexicaul, uppermost leaves (on peduncles) scaly, indistinguishable 
from outer involucral bracts. Capitula with about 10 florets, almost 
cylindrical or turbinate, with fruits 7-10 mm long, solitary at apices of 
long slender peduncles. Involucral bracts mostly three-rowed; outer bracts 
narrowly triangular, ovate; inner bracts longer, linear-lanceolate, with 
membranous border, obtuse, usually dark violet and with barbula of fine 
light-colored hairs (under a hand lens!), in remaining part dorsally green, 
very short appressed-hairy with papilliform hairs (under high 
magnification!). Corolla lilac-colored(?). Achenes about 5 mm long and 
1 mm wide, linear-elliptical, light yellow, slightly flattened, with rather 
numerous, but not always prominent longitudinal ribs, of which only three 
or four distinct, covered with short, light-colored hairs (indistinct even 
under high magnification), apically attenuate, bearing thin slightly distinct, 
about 1 mm long beak; inner row of pappus hairs firmly attched, 4-5 
mm long. Flowering June to July. (Plate XX, Fig. 8.) 

At 3,000-3,500 m, along banks of mountain streams and lakes or 
near edges of glaciers; less often in middle mountain zone in juniper 
woodlands —Soviet Central Asia: mountainous Turkmenia (Chopan-Dag), 
Pamiro-Alai (Darvaz), mountainous Badakhshan. General distribution: Iran 
Region (Iraq, Afghanistan). Described from Iran. Type in Geneva; isotype 
(collections of Biinge) in Leningrad. 

Note. The species is sharply isolated from the remaining members of 



347 

the genus Cephalorrhynchus and is being included by us in this genus 
only on the basis of the formal character of the achene structure. The 
nature of achene as well as other morphological data rule out the possibility 
of retaining this species in the genus Cicerbita. It also cannot be left in 
the genus Lactuca, as initially proposed by Krascheninnikov, because the 
presence of an outer series of pappus is not a typical characteristic of 
Lactuca species. Either this unique species should be treated as a separate 
genus, or it should be put, as a separate section, in Cephalorrhynchus, to 
which it is close on the basis of achene structure, which, for the present, 
we have done. 

GENUS 1659. Cicerbita Wallr. '^ 

Wallr. Sched. crit. I (1822) A^?).— Cicerbita Wallr. emend. Beauverd 
in Bull. Soc. Bot. Geneva, 2 ser. II (1910) 115, p. min. p. (seorsum 
impressum: Contrib. a I'etude des Compos. Ill, p. 17). — Agathyrsus D. 
Don in Edinb. New Phil. Joum. VI (1829) 310, p. p.—Mulgedium sect. 
Eumulgedium DC. Prodr. VII, I (1838) 248, p. p.— Lactuca sect. 4 
Cicerbita Benth. in Benth. and Hook. f. Gen. pi. II (1873) 525, p. p. 
352 Capitula homogamous, with 10-25(30)florets or (less often) 5-8- 

florets, small (fruits reaching 10-15 nmi in length), mostly cylindrical, in 
racemose, racemose-paniculate, or corymbose inflorescence. Involucre 
usually two- or three-rowed; involucral bracts herbaceous, dorsally finely 
appressed-hairy or densly covered with short papillae (under a hand lens!), 
often mixed with glandular hairs; innermost bracts two to four times as 
long as outermost, with membranous border, sometimes with beard of 
fine hairs at apex (under a hand lens!). Form of achenes distinctly variable: 
oblong, not or only slightly flattened, or lanceolate to ellipsoid (narrowly 
linear- or oblong-ellipsoid), more or less distinctly flattened to almost 
flat, somefimes flattened-triangular; achenes with very short, sometimes 
with deep constriction just below apex, or apex narrowed into filiform 
structure, but in both cases achene terminating in a broad (almost equaling 
its broadest part), somewhat flat disk bearing pappus, more or less densely 
covered throughout with very short hairs, less often almost glabrous; 
pappus of outer row of hairs strongly attached to disk, coronate (under a 
hand lens!) and inner row consisting of somewhat longer, thin, white or 
dirty-white hairs, usually falling easily, usually appearing smooth (under 
high magnification, barbed). 



' Treatment by M.E. Kirpicznikov. 

^ Name of the plant found in the work of Pope Marcellinus (4th century AD) and 
corresponds to the name Sonchus. 



348 

Lectotype of genus: Cicerbita alpina (L.) Wallr. 

Perennial, mainly rhizomatous herbs, usually with solitary and erect 
stems, within the USSR found in mountain regions of the Caucasus and 
Soviet Central Asia, much less frequently in the Carpathians, as well as 
in the mountains of the Altai in Siberia. Plants primarily growing in forests, 
in shrub thickets and subalpine meadows. 

The number of species in this genus, as well as its characteristics, 
are quite uncertain. It is not possible to agree with Beauverd (I.e.), who 
interpreted the genus Cicerbita in an extremely broad sense, including 
under it Mulgedium Cass., Lactucopsis Sch. Bip., Mycelis Cass., 
Cephalorrhynchus Boiss., and Steptorhamphus Bge. In this sense, the 
genus Cicerbita included more than 50 species already in 1910. On 
the other hand, the well-known specialist on the Cichorieae, Ledyard 
Stebbins (Critical notes on Lactuca and related genera. Journ. of Botany, 
London, LXXV, 1937), proposed leaving only two species, C. alpina 
(L.) Wallr. and C. pancicii (Vis.) Beauverd, in the genus Cicerbita and 
including in Lactuca the genera Mulgedium, Lactucopsis, Mycelis, 
Scariola and a large part of Cicerbita in the sense of Beauverd. It is 
difficult also to agree fully with this viewpoint. For example, it is 
unthinkable to countenance Lactuca sativa near Cicerbita zeravschanica, 
because they differ in almost all characters, from plant habit to achene 
structure. 

We adopt, tentatively, for the genus Cicerbita, fairly broad limits, 
353 being convinced, however, that further study will require raising the 
subgenera of this genus (and maybe even certain sections) to the level of 
separate genera. 

1. Florets in capitulum usually five or six, less often eight; stem in 
upper part covered with squamiform glandular hairs 2. 

+ Florets in capitulum 10 or more, very rarely fewer, but then stem 
in upper part without squamiform hairs 4. 

2. Corolla yellow; basal leaves more or less densely covered with 
narrow squamiform hairs beneath 

11. C. zeravschanica M. Pop. ex S. Kovalevsk. 

+ Corolla lilac or pink 3. 

3. Middle cauline leaves elliptical, often large (10-15 cm long and 
7-9 cm wide); capitula usually with five-florets, with pappus hairs 
not exceeding involucre; inflorescence a lax panicle. Species 
endemic to eastern Fergana 13. C. kovalevskiana Kirp. 

+ Middle cauline leaves much smaller, with laminas often more or 
less incised; capitula with five- or six (eight) florets, with pappus 
hairs slightly exceeding involucre; inflorescence corymbose- 
paniculate or almost corymbose. Species primarily growing in the 



349 

western part of Tien Shan 

12. C. rosea (M. Pop. and Vved.) S. Kovalevsk. 

4(1) Achenes oblong, not flattened or almost not flattened, dark brown, 
more or less glabrous or covered with extremely short hairs (under 
high magnification!) with very short, often deep constriction at 

apex; outer corona of pappus hairs not always distinct 5. 

+ Achenes lanceolate or ellipsoid, more or less flattened to almost 
completely flat or flattened-triangular, usually dark violet, less often 
olive-colored, narrowed throat-like at apex 7. 

5. Involucral bracts dorsally covered with more or less numerous, 
squamiform, glandular hairs; stem glandular-hairy above 6. 

+. Involucral bracts finely appressed-hairy dorsally (under a lens!); 
stem glabrous or subglabrous. Relatively robust plant (100-125 
cm high), with thick woody undergrounnd part, within the USSR 

growing only in Soviet Central Asia 

2. C. thianschanica (Rgl. and Schmalh.) Beauverd. 

6. Rhizome up to 10 cm long and 1 cm thick; stem in lower part 
covered with narrow, squamiform hairs; pappus hairs mostly dirty 
white, less often white, about two times as long as achene. Species 
found in the north of the European part of the USSR and in the 
Carpathians 1. C. alpine (L.) Wallr. 

+ Rhizome short, 2-5 mm thick at base; stem in lower part glabrous; 
pappus hairs snow-white, about one and one-half times as long as 

354 achene. Species found in Siberia and Soviet Central Asia 

3. C. azurea (Ldb.) Beauverd 

7(4) Corolla yellow; outer pappus hairs particularly distinct; achene apex 

attenuate into relatively narrow, throat-like structure 

10 C. deltoide (M.B.) Beauverd 

-I- Corolla light blue, lilac, or azure; outer pappus hairs more or less 
distinct; achene apex attenuate into broad, throat-like structure .. 
8. 

8. Plant robust, herbaceous, up to 8-200 (250) cm high; stem 10-15 
mm in thick at base; capitula 25-30-flowered, broadly cylindrical; 
peduncles densely glandular-hairy 9. 

-I- Plant shorter or stem slender at base, capitula usually with fewer 
flowers, or peduncles not densely glandular-hairy 10. 

9. Inflorescence branches 1.5-2(3) mm wide at base, shghtly arcuately 
curved, lateral branches more or less distinctly exceeding central 
axis; achenes olive-colored; inner pappus hairs white. Plants 
growing in the European part of the USSR and in Western Siberia 

4. C. uralensis (Rouy) Beauverd 

-I- Inflorescence branches relatively thick, largest branches 3-5 mm 
thick, usually arcuately curved upward, lateral branches mostly 



350 

distinctly exceeding central axis; achenes dark violet; inner pappus 

hairs dirty white. Plants growing in the Caucasus 

5. C. macrophylla (WiUd.) Wall. 

10(8) Achenes, at least in one plane, with only one middle well 
developed longitudinal rib 11. 

+ Achenes with more or less numerous, almost equally prominent 
longitudinal ribs 12. 

11 Stem in upper part, as also involucral bracts dorsally, covered 
with squamiform, more or less glandular hairs; inner pappus hairs 
usually dirty white 6. C. sevanensis Kirp. 

+ Stem in upper part glabrous or finely acachnoid-tomentose; 
involucral bracts dorsally covered with dense, short, papillose hairs 

(under a hand lens!), eglandular 

7. C. racemosa (Willd.) Beauverd. 

12(10) Outer row of pappus hardly distinguishable; disk densly covered 
with very thin arachnoid hairs (under a hand lens!); inflorescence 

often up to 30-50 cm or longer 

9. C. bourgaei (Boiss.) Beauverd 

+ Outer row of pappus distinguishable (under a hand lens!); disk 
not covered with thin acachnoid hairs; inflorescence usually less 
robust 8. C. prenanthoides (M.B.) Beauverd. 

355 Subgenus 1. Cicerbita. — Sonchus sect. I. Cicerbitae (Wallr.) W.D. 

Koch, Synops. fl. Germ. et. Helvet. (1837) 433. — Mulgedium (sect.) II. 
Cicerbita C.A. Mey. in Mat. k BUzh. Pozn. Prozyab. Ross. Imp. VI (1849) 
26 (Verzeichn. Dr. Kolenati gesammelt. Pflanz.). — Achenes oblong, not 
flattened or only slightly compressed, dark brown, with more or less 
numerous longitudinal ribs, some of them slightly more prominent, almost 
glabrous or covered with extremely short hairs (under high magnification!), 
slightly narrowed toward base, with very short, sometimes rather deep 
constriction at apex, terminating in broad disk bearing pappus; outer row 
of pappus (under a hand lens!) not always distinct, inner comprising 
usually rather easily falling hairs. 

Type of subgenus: type of genus. 

Note. Achenes, if outer whorl of pappus is not considered, are fairly 
similar to achenes of typical Prenanthes. 

1. C. alpina (L.) Wallr. Sched. crit. I (1822) 434; Beauverd in Bull. 
Soc. Bot. Geneve, 2 Ser. II, 118 (seorsum impressum: Contrib. a I'etude 
des Compos. Ill, p. 20, No. 1); Klokov in Vizn. Rosl. URSR 602; Hayek 
in Hegi, 111. Fl. 2 Aufl. VI, 2 1099; Stank, and Tal. Opred. Vyssh. Rast. 
(1957) 442. — Sonchus alpinus L. Sp. pi. (1753) 794. — Hieracium 
coeruleum Scop. Fl. camiol. ed. 2, II (1772) 111. — Sonchus montanus 
Lam. Encycl. meth. Ill (1791-1792) 401.— ?5. coeruleus Smith, Fl. Brit. 



351 

II (1804) 815. — lAracium alpinum Monn. Essai Monogr. Hierac. (1829) 
73. — Mulgedium alpinum Cass, ex Less. Synops. Compos. (1832) 142; 
DC. Prodr. VII, 1, 248; Lbd. Fl. Ross. II, 2, 840; Mishkin, Fl. Khibinsk. 
Gor. 70. — Lactuca alpine A. Gray, Synoptic. Fl. N. Americ. I, 2 (1884) 
444.— Ic: RchK f. Ic. fl. Germ, et Helvet, XIX (1859), t. 64 (MCCCCXV) 
(sub Mulgedio alpino); Javorka and Csapody, Iconogr. fl. Hungar. (1933), 
t. 559, fig. 4017, Hegi, op. cit. t. 277, fig. 2 (color.).— Exs.: Rchb. Fl. 
Germ. exs. 1820, No. 2427 (sub Mulgedio alpino (L.) Walk.); Billot, Fl. 
Gall, et Germ. exs. No. 2104 (sub M. alpino L.); F. Schultz, Herb. norm, 
cent. 17, No. 1602 (sub M. alpino Less.); Hayek, Fl. Stir. exs. No. 493 
(sub M. alpino Cass, in Less.); Petrak, Fl. Bohem, et Morav. exs. No. 
1096 (sub nomine eodem); Fl. exs. reip. Bohem. Slov. No. 277 (ut supra); 
PI. Finl. exs. No. 1410 (sub Lactuca alpina (L.) A. Gray); Fl. Bulg. exs. 
(sine num., sub Mulgedio alpino Cass.); Sam. PI. Suec. exs. ed. Hult. No. 
1609 (sub nom. paenultimo). 

Perennial. Rhizome up to 10 cm long and about 1 cm thick. Stem 
50-100 cm (250 cm) high, sulcate, fistular, in lower part covered with 
narrow, squamiform hairs, glabrous or almost so above, but densely 
covered with narrow, squamiform, glandular hairs above; hairs often dark 
brown red or violet. Leaves thin, green above, glaucous beneath; lower 
and middle cauline leaves lyrate or lyrate-runcinately divided, with large 
terminal triangular or triangular-hastate segment, and winged, auriculate, 
356 semiamplexicaul petiole, sinuate-toothed, along margin with soft spinules; 
apical leaves strongly reduced, lanceolate to triangular and narrowly linear, 
scaly at base and especially on peduncles. Capitula mostly with 15-20- 
florets obtruncate-conical or almost cylindrical, with fruits 12-15 mm 
long, in racemose or narrowly racemose-paniculate inflorescence at apices 
of slender, densely glandular-hairy peduncles. Involucral bracts mostly 
two-rowed, dorsally covered with squamiform and usually glandular 
hairs, often violet. Corolla light blue-violet or yellow. Achenes (3)3.5- 
4(5) mm long and about 1 mm wide, oblong, sometimes slightly 
compressed, light brown, with numerous strongly projecting longitudinal 
ribs, finely puberulent (under high magnification!) or almost glabrous, 
slightly narrowed at base, with rather short and shallow constriction at 
apex, terminating in broad disk bearing pappus; outer pappus hairs very 
short, dense, forming corona (under a hand lens!); inner hairs falling rather 
easily, unequal, but nearly two times as long as achene, dirty white or 
white (often yellow at base). Flowering June to July (September). (Plate 
XXI, Fig. 2.) 

On Kola Peninsula in spruce and birch forests, depressions, spring 
marshes, banks of brooks, less often on meadow-covered slopes; in the 
Carpathians in beech and spruce forests, at relatively high altitudes (reliable 
reports for the USSR up to 1,200 m, and in the mountains of Europe, up 



352 




357 Plate XXII. 

1, 5 — Cicerbita kovalevskiana Kirp., habit of plant, achene; 2 — C. deltoidea (M.B.) 

Beauverd, achene; 3 — C. zeravschanica M.Pop. ex S. Kovalevsk., achene; A — C. rosea (M. 

Pop. and Vved.) S. Kovalevsk., achene. 



353 

to 2,200 m). — Arctic: Arctic Europe (Rybachii Peninsula, Kildin Island 
and northern part of Kola Peninsula); European Part: Karelia-Lapland 
(Kola Peninsula), Upper Dniester (Carpathians). General distribution: 
Scandinavia, central and Atlantic Europe, western Mediterranean, Balkans- 
Asia Minor (northern Balkan Peninsula). Described from Europe (from 
the high mountains of Lapland, Switzerland and Austria). Type in London. 
Note. Ledebour (1. c. 841), apparently for no good reason, included 
"Sonchus canadensis With." as a synonym of this species. 

2. C. thianschanica (Rgl. and Schmalh.) Beauverd in Bull. Soc. Bot. 
Geneve, 2 Ser. II (1910) 123 (seorsum impressum: Contrib. a 1' etude des 
Compos. Ill, p. 25, No. 12). — Mulgedium thianschanicum Rgl. and 
Schmalh. in Tr. Peterb. Bot. Sada VI, 2 (1880) 329; O. and B. Fedtsch. 
Perech. Rast. Turk. IV, 350 (Consp. Fl. Turk. No. 2855). 

Perennial. Rhizome thick, woody. Stem 100-125 cm high, 10-15 
mm thick at base, sulcate, fistular, glabrous or subglabrous. Leaves thin, 
unequally toothed, teeth softly cuspidate; lower leaves on long, winged 
358 petiole, semiamplexicaul, large (often 40-50 cm long and 7-9 cm wide), 
lyrately pinnatisect, with large terminal, triangular-hastate segment and 
few small lateral segments; middle cauline leaves sessile, more or less 
like lower leaves in form; upper leaves smaller, with broad, almost 
amplexicaul base and more or less undivided, oblong or lanceolate lamina; 
leaves on peduncles scaly. Capitula usually with 15-20 florets, cylindrical 
or slightly broadened above, with fruits about 15 mm long and 7-8 mm 
wide, borne in narrow but long (30-50 cm and longer) racemose 
inflorescence; often inflorescence branches arising from nodes in clusters 
(few at a time). Involucral bracts mostly three-rowed, somewhat lax, 
dorsally finely appressed-hairy (under a hand lens!), at least tip usually 
violet. Corolla light blue. Achenes about 6 mm long and slightly more 
than 1 mm wide, oblong, sometimes slightly compressed, brown or dark 
brown, with more or less numerous logitudinal ribs covered with very 
short, thin, light colored, upward-directed hairs (under high magnification!) 
or almost glabrous, apically achenes more or less deeply constricted, 
terminating in four- or five-angled roundish disk bearing a pappus; outer 
hairs of pappus very short (about 0.4 mm long), numerous, dense, forming 
corona; inner hairs fragile, white, easily falling, not strictly equal, 6-7 
mm long. Flowering June to September. (Plate XXI, Fig.l.) 

At altitudes from 1,500 to 3,500 m; in spruce forests of Picea 
schrenkiana; among trees, scrubs, in herb meadows and on grassy slopes; 
floodplains and river banks; near glaciers. — Soviet Central Asia: 
Dzhungaria-Tarbagatai (Dzhungaria), Syr-Darya (one doubtful specimen 
from Turpak-Bel), Pamiro-Alai, Tien Shan. General distribution: 
Dzhungaria-Kashgaria (Dzhungaria). Described by Regel from the Muzart 
Valley. Type and isotype in Leningrad. 



354 

3. C. azurea (Ldb.) Beauverd in Bull. Soc. Bot. Geneve, 2 Ser. II 
(1910) 123 (seorsum impressum: Contrib. a I'etude des Compos. Ill, p 
25, No. 10); Kryl. Fl. Zap. Sib. XI, 3021; Sobolevskaya, Konsp. Fl. Tuvy 
200; Grubov, Konsp. Fl. MNR, 282, No. 1855 (Tr. Mong. Kom. Vyp 
67). — Sonchus azureus Lbd. Fl. Alt. IV (1833) 138; Ej. Icon. pi. fl. Ross 
V (1834) 21.—Mulgedium azureum DC. Prodr. VII, 1 (1838) 248; Ldb 
Fl. Ross II, 2, 842; O. and B. Fedtsch. Perech. Rast. Turk. IV, 350 (Consp 
Fl. Turk. No. 2856); Popov, Fl. Sr. Sib. II, 786; Nikitina, Mater, po Fl 
Sev. Ski. Khr. Kirg. Ala-too, 110, No. 1125. — Lactuca azurea Danguy in 
Bull. Mus. Nat. Hist. Paris XX (1914) 39; Ling in Contrib. Inst. Bot. 
Nat. Akad. Peip. Ill, 4, 185. — Ic: Ldb. op. cit. t. 496 (color., bona). — 
Exs.: Kar. and Kir. 1840, No. 895; Smim. PI. alt. exs. No. 98. 

Perennial. Rhizome slender, 2-5 mm thick at base, short. Stem 
(35)50-70 cm high, slightly sinuate, glabrous below, weakly branched 
359 above and more or less densely covered with narrow, squamiform, usually 
glandular hairs. Leaves thin, well developed, mainly crowded in lower 
part of stem, green above, gray or glaucous beneath, tomentose, sometimes 
also with scattered, narrow, squamiform hairs (mainly on veins); lower 
leaves on narrow-winged, more or less amplexicaul petioles expanded at 
base, two times or more as long as cordate- or ovate-hastate or reniform 
lamina; usually lamina more or less lyrately pinnatisect, and then with 
one to many small lateral segments; middle cauline leaves smaller than 
lower, hastate-rhombic or triangular, on shorter and more or less broad- 
winged petioles; uppermost leaves linear-lanceolate or lanceolate, reduced, 
in inflorrescence scaly. Capitula with 10-12(18) florets, with fruits almost 
cylindrical, 10-13 mm long, on peduncles densely covered with narrow, 
squamiform, glandular hairs, forming lax racemose (less often racemose- 
paniculate or paniculate-corymbose) inflorescence. Involucral bracts mostly 
two-rowed, usually blackish-violet, with more or less numerous glandular 
hairs on dorsal surface. Corolla bright azure or dark blue. Achenes about 
4 mm long and 1 mm wide, oblong, sometimes slightly compressed, with 
numerous prominent longitudinal ribs, dark brown, puberulent with light- 
colored hairs (under high magnification), apically with short, more or 
less deep constriction, terminating in broad, roundish, four- or five-angled 
disk, bearing pappus; outer hairs of pappus very short (sometimes 
indistinct), forming corona, snow-white, easily falling, about 6 mm long. 
Flowering June to August(September). (Plate XXI, Fig. 3.) 

Deciduous and cedar forests, up to their upper limits in subalpine 
and alpine meadows (Siberia, Mongolia), spruce forests of Picea 
schrenkiana (Soviet Central Asia). — Western Siberia: Altai; Eastern 
Siberia: Angara-Sayans; Soviet Central Asia: Dzhungaria-Tarbagatai, Tien 
Shan. General distribution: Dzhungaria-Kashgaria, Mongolia. Described 
from Altai. Type in Leningrad. 



355 

Subgenus 2. Platyachena Kirp. subgen. nov. in Addenda XXVIII, 
726. — Achenes lanceolate or linear-ellipsoid, somewhat compressed to flat 
or flattered-triangular, usually dark violet, with more or less numerous, 
less often one to few ribs on each side, entirely puberulent, with hairs in 
transversely sinuate rows; achenes apically constricted in throat-like 
structures. 

Type of subgenus: Cicerbita uralensis (Rouy) Beauverd. 

Section 1. Platyachena Kirp. sect, nova in Addenda XXVIII, 726. — 
Achenes more or less compressed or flattened-traingular, at least in one 
360 plane with only median distinctly rib, entirely puberulent with hairs in 
tranversely sinuate rows (under a hand lens!). 

Type of section: type of subgenus. 

4. C. uralensis (Rouy) Beauverd in Bull. Soc. Bot. Geneva, 2 ser. II 
(1910) 123 (seorsum impressum: Contrib. a I'etude des Compos. Ill, p. 
25, No. 14). — Mulgedium uralense Rouy, Illustr. pi. Eur. rar. fasc. 16 
(1901) t. 390 (non widil). —Sonchus hispidus Ldb. Fl. Alt. IV (1833) 140, 
in adnot. — Mulgedium hispidum DC. Prodr. VII, 1 (1838) 250; Ldb. Fl. 
Ross. II, 2, 844; Kraschen, in Fl. Yugo-Vost. VI, 465; Kirp. in Majevski, 
Fl. (1954) 629. — M. macrophyllum DC. var. hispidum Korsh. in Mem. 
Acad. Sc. Petersb. VIII, ser. VII, 1 (1898) 265 (Tentam. fl. Ross. or. No. 
778). — Cicerbita gmelinii Beauverd, op. cit. (seorsum impressum: 1. c. 
No. 11); Kryl. Fl. Zap. Sib. XI, 3021; Stank, and Tal. Opred. Vyssh. 
Rast. (1957) 442. — Mulgedium macrophyllum auct. non DC. (incl. var. 
hispidum Korsh.); Fedtsch. and Fler. Fl. Evrop. Ross. 1055; Govorukhin, 
Fl. Urala 521.— Ic: J.G. Gmel. Fl. Sibir. II (1749) t. IV fig. 1. 

Perennial. Rhizome short, densely covered with tuft of filiform thin 
dark-brown roots. Stem 80-200 cm high, up to 10-15 mm thick at base, 
sulcate, fistular, usually glabrous or subglabrous below, more or less 
densely covered with narrow (almost filiform), squamiform, glandular hairs 
above, especially from point of branching. Leaves thin, mostly with very 
narrow, pale, squamiform hairs beneath as well as along margin; lowermost 
leaves large (including petiole up to 15 cm long), hastate- or cordate- 
triangular, on long, broadly winged, semiamplexicaul petiole, often divided 
and then with one to many small, lateral segments; middle cauline leaves 
lyrate, with large, hastate, apical segment and often also with one pair of 
small lateral segments; upper leaves smaller, usually sessile, almost 
amplexicaul, oblong-lanceolate or lanceolate, often unevenly toothed, 
sometimes sinuate-spatulate, with numerous, very narrow, squamiform, 
glandular hairs beneath; uppermost leaves reduced; leaves on peduncles 
hardly distinguishable from involucral bracts. Capitula usually with 25- 
30 florets, broadly cylindrical, with fruits (10)14-15 (17) mm long, and 6- 



356 

10 mm wide, in broad and often long paniculate-corymbose inflorescence; 
inflorescence branches 1.5-2(3) mm thick at base, more or less arcuately 
curved, lateral branches more or less distinctly exceeding central axis, all 
branches covered with narrow, squamiform, glandular hairs, especially 
dense on peduncles bearing capitula. Involucral bracts mostly three-rowed, 
dorsally with short, appressed, papilliform hairs (under high 
magnification!) and more or less dense glandular hairs. Corolla light blue. 
Achenes about 6 mm long and 1.5 nrni wide, lanceolate or linear-ellipsoid, 
compressed, olive-colored when mature, with one prominent midrib and 

361 often two or three additional, more or less distinct, longitudinal ribs on 
each side, with throat-like constriction at apex; outer row of pappus hairs 
very short (under a hand lens!), inner row of easily faiUng, fragile, white, 
7-8 mm long hairs. Flowering June to September. (Plate XXI, Fig. 4.) 

Shady moist places; broad-leaved, coniferous, and mixed forests, forest 
edges, forest glades and logging areas in thickets of tall grass; alder 
thickets, banks of streams and in forest ravines; subalpine herb meadows 
to 1,000 m. — European Part: Upper Volga, Volga-Kama, Volga-Don 
(northern part; based on literature records). Trans- Volga Region; Western 
Siberia; ?0b (extreme west). Upper Tobol (western part). Endemic. 
Described from Verkhotur (Sverdlovsk Region) at the border of Volga- 
Kama and Ob regions of the "Flora of the USSR"). Type in Leningrad. 

Note. This species is very close to Cicerbita macrophylla (Willd.) 
Wallr. and morphologically quite difficult to distinguish from it. In spite 
of good geographic isolation, the separation of C. macrophylla and C. 
uralensis at the specific level is in need of careful study. 

5. C. macrophylla (Willd.) Walk. Sched. crit. I (1822) 434; Beauverd 
in Bull Soc. Bot. Geneve, 2 Ser. II, 124 (seorsum impressum): Contrib. a 
I'etude des Compos. Ill, p. 26. No. 16); Grossh. Fl. Kavk. IV (1934) 253 
— Sonchus macrophyllus Willd. Sp. pi. Ill; 3 (1803) 1519. — Mulgedium 
macrophyllum DC. Prodr. VII, 1 (1838) 248; Boiss. Fl. or. Ill (1875) 799 
(incl. p. glabrum Boiss.). — M. grande C. Koch in Linnaea XVII (1843) 
279 (incl p. glabrum C. Koch); Lbd. Fl. Ross. II, 2, 841.— M glabrum 
C. Koch in Linnaea XXIII (1850) 668. — Lactuca macrophylla A. Gray, 
Synoptic. Fl. N. Americ. I, 2 (1884) 444.— L. macrophylla Schmalh. Fl. 

11 (1897) 166. — Mulgedium acuminatum Conrath and Freyn ex Freyn in 
Bull. Herb. Boiss. Ill, 9 (1895) 476 (non M. acuminatum DC. Prodr. VII, 
1 (1838) 250). — Cicerbita conrathiana Beauverd, op. cit. p. 125 (seorsum 
impressum: p. 27, No. 19). — C acuminata (Conrath and Freyn ex Freyn) 
Grossh. Fl. Kavk. IV (1934) 253 (nee C. acuminata Walk. Sched. crit. I 
(1822) 434). — C. grandis (C. Koch) Schchian in Zam. po. Sist. i Geogr. 
Rast. Tbil. 13 (1947) 89 and in H. Gruzii VIII, 673; Grossh. Opred. 
Rast. Kavk. 515; Askerova in Fl. Azerb. VIII, 546. 



357 

Perennial. Rhizome short, densely covered with tuft of slender filiform 
dark brown roots. Stem (90)150-200(250) cm high, up to 10-15 mm 
thick at base, sulcate, fistular, mostly hollow, glabrous or almost glabrous 
below, above, particularly from point of branching, more or less densely 
covered with thin squamiform, glandular hairs (only very rarely 
inflorescence branches glabrous or almost glabrous — var. glabra (C. Koch) 
Kirp. comb. nova. h. 1.; basionymum: Mulgedium glabrum C. Koch I.e.). 
Leaves thin, almost always covered with thin, squamiform, often glandular, 
hairs beneath along main and lateral veins and also along margin; lower 
leaves large (up to 40 cm long including petiole), ovate- 
362 or hastate-triangular, less often broadly rotund-ovate, on long, broadly 
winged, semiamplexicaul petiole, often also with one or two pairs of small 
lateral segments; middle cauline leaves rather large (10-20 cm long), 
broadly lanceolate or oblong-ovate, with broad semiamplexicaul base, and 
well developed cusp at apex; upper cauline leaves smaller, sessile or 
subsessile, semiamplexicaul, highly variable (mostly oblong-lanceolate, 
oblong or linear-lanceolate, less often panduriform or other shapes), usually 
with numerous, squamiform, glandular hairs beneath; uppermost leaves 
reduced, on peduncles hardly distinguishable from involucral bracts. 
Capitula usually with 25 florets, broadly cylindrical or narrow- 
campanulate, with fruits 12-14(16) mm long, in paniculate-corymbose 
inflorescence, often large (30-50 cm long, sometimes longer); largest 
inflorescence branches 3-5 mm thick at base, usually arcuately curved 
upward, lateral branches usually distinctly exceeding central axis, all 
covered with scabrous, thin, squamiform, glandular hairs, especially 
densely on peduncles bearing capitula. Involucral bracts mostly three- 
rowed, with short, appressed, papillose pubescence on dorsal surface (under 
high magnification!) and more or less numerous, thin, squamiform, 
glandular hairs. Corolla light blue. Achenes about 6 mm long and 1.5 
mm (sometimes 2 mm) wide, lanceolate or linear-ellipsoid, somewhat 
flat, dark violet at maturity, with one or two prominent longitudinal ribs 
on each side, with throat-like constriction at apex; outer row of pappus 
hairs very short, dense, sometimes yellowish, inner hairs fragile, easily 
falling, usually dirty-white, 6-7.5 mm long. Flowering June to August. 
(Plate XXI, Fig. 5) 

Montane forests, shrub thickets and subalpine meadows, in more or 
less shady places. — Caucasus: all regions except Talysh. Endemic (?). 
(Possibly growing in Turkey.) The native region (America) in the original 
description is in error. Type in Berlin. 

Note. Morphologically, this species is quite similar to C. uralensis. 

6. C. sevanensis Kirp. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk 
SSSR XXII (1963) 304. 



358 

Perennial. Rhizome rather long (5 cm or more), about 1 cm thick, 
covered with brown remnants of dead leaves. Stem 70-100 cm high, 5- 
8 nmi thick at base, sulcate, fistular or more or less solid, covered with 
thin, squamiform, and more or less glandular hairs, usually dense only 
above, especially from point of branching; glabrous or almost glabrous 
in middle. Leaves thin, semiamplexicaul, with narrow, squamiform, 
363 glandular hairs along veins and margin, or such hairs almost absent; lower 
leaves rather large (up to 20 cm long including petiole), mostly lyrate, 
with broadly ovate or hastate-triangular terminal segment and broad- 
winged petiole, strongly expanded like sheath at base, sometimes also 
with one pair of small lateral segments, middle cauline leaves usually 
broadly oblong-ovate or broadly lanceolate, with semiamplexicaul base, 
and well-developed cusp at apex; upper cauline leaves oblong-ovate, 
lanceolate or narrowly triangular, mostly very finely appressed tomentose- 
arachnoid beneath (under a hand lens!) mixed with thin, squamiform, 
glandular hairs, especially along midrib; uppermost leaves strongly 
reduced, on peduncles hardly distinguishable from involucral bracts. 
Capitula mostly with 25 florets, broadly cylindrical or campanulate, with 
fruits 13-15 mm long, borne in not very large paniculate-corymbose 
inflorescence (mostly 10-25 cm long); largest branches of inflorescence 
not exceeding 3 mm in thickness, straight or only slightly arcuate, more 
or less densely covered with scabrous, thin, squamiform hairs, especially 
dense on peduncles bearing capitula. Involucral bracts mostly three-rowed, 
with short appressed hairs on dorsal surface (under high magnification!) 
and more or less dense, thin, glandular, squamiform hairs. Corolla light 
blue. Achenes about 5 mm long and 1.5 nun wide, lanceolate or linear- 
ellipsoid, compressed or flat-trihedral, mature achenes dark violet, with 
one or two (sometimes more) prominent longitudinal ribs on each side, 
and throat-like constriction at apex; pappus hairs of outer row very short, 
dense, usually dirty white; inner hairs fragile, easily falUng, usually white, 
almost entirely smooth, up to 7-8 mm long. Flowering July to August. 
(Plate XXI, Fig. 6). 

Montane forests and subalpine meadows. — Caucasus: ?Western 
Transcaucasia, Eastern and Southern Transcaucasia. Endemic. Described 
from vicinity of Lake Sevan. Type and isotype in Leningrad. 

7. C. racemosa (Willd.) Beauverd in Bull. Soc. Bot. Geneve, 2 Ser. 
II (1910) 122 (seorsum impressum: Contrib. a I'etude des Compos. Ill, p. 
24, no. 8); Grossh. Fl. Kavk. IV (1934) 253 and Opred. Rast. Kavk. 514; 
Kolak. Fl. Abkhazii IV, 299; Schchian in Fl. Gruzii VIII, 673; Askerova 
in Fl. Azerb. VIII, 546.— Lactuca racemosa Willd. Sp. pi. Ill, 3 (1803) 
l534.—Sonchus sibiricus M.B. Fl. taur.- cauc. II (1808) 240 (non 5. 
sibiricus L.). — Sonchus albanus Stev. in Mem. Soc. Nat. Mosc. IV (1813) 



359 

99. — Lactuca albana C.A. Mey. Verzeichn. Pfl. Cauc. (1831) 56. — 
Mulgedium albanum DC. Prodr. VII, 1 (1838) 249; Ldb. Fl. Ross. II, 2, 
844; Boiss. Fl. or. Ill, 800. — M. albanum DC. var. glaberrima Trautv. 
and var. setulosa Trautv. in Tr. Peterb. Bot. Sada, IV, 2 (1876) 386.— 
Lactuca albana Schmalh. Fl. II (1897) 166. — Mulgedium racemosum 
(Willd.) Schischk. in Grossh. and Schischk. Sched. ad herb. "PI. Orient 
exsiccatae," fasc. I-VIII (1924) 44.— Exs.: Fl. Cauc. exs. No. 250 (sub 
364 Mulgedio albano DC. var. glaberrimo Trautv.) PI. or. exs. No. 175 (sub M. 
racemoso Schischk.); Sintenis, It. or. 1894, No. 6182 (sub M. albano DC). 
Perennial. Rhizome slender, producing long funiform roots. Stem 
(10)30-60(110) cm high, slightly sulcate, almost glabrous or unevenly 
and finely arachnoid-tomentose, very rarely covered with thin squamiform 
hairs at base, branched above or just near tip (sometimes almost from 
base). Leaves (well developed!) usually few, crowded in lower part of 
stem; lower leaves more or less lyrate-runcinate or lyrate, basally narrowed 
into winged petiole; other leaves entire or subentire to incised to various 
degrees, on winged petiole or sessile, semiamplexicaul, like lower leaves 
sinuate-toothed or sinuate-spinose, sometimes almost serrate-spinose, more 
or less cuspidate, glabrous beneath, appressed-hairy or with thin 
squamiform hairs; upper cauline leaves few, oblong and lanceolate to 
narrow-triangular and linear, uppermost leaves scaly. Capitula with about 
20 florets, almost cylindrical, with fruits 12-14 mm long, usually 
assembled in few-headed corymbose panicle or panicle; peduncles slender, 
more or less arachnoid-pubescent or occasionally covered with thin 
squamiform hairs (var. setulosa (Trautv.) Kirp. comb, nova h. 1.; 
basionymum — Mulgedium albanum var. setulosa Trautv. 1. c). Involucral 
bracts mostly three-rowed, with dense, short, papillose hairs on dorsal 
surface (under high magnification!); inner bracts with barbula of very 
short, thin hairs at apex (under a hand lens!); corolla light blue or lilac. 
Achenes about 5 mm long and slightly more than 1 mm wide, lanceolate 
or narrow-ellipsoid, somewhat flat, mature achenes dark violet, with one 
to many prominent longitudinal (sometimes winged) ribs, with throat-like 
constriction at apex; pappus two-rowed, hairs of outer row dense, often 
yellowish, short (under a hand lens!) forming a corona; hairs of inner 
row snow-white, 6-7 mm long. Flowering (May) June to August 
(September). (Plate XXI, Fig. 7.) 

More or less shady places in montane forests and subalpine meadows, 
on stony slopes and screes, in glacial moraines, sometimes in cultivated 
fields; at aUitudes of (800)1,300-3,000 m. — Caucasus: Ciscaucasia, 
Dagestan, Eastern and Western Transcaucasia, Southern Transcaucasia 
(rarely?). General distribution: Balkans-Asia Minor, Armenia and 
Kurdistan. Described by Willdenow from "Armenia" ("Habitat in 
Armenia"). Type in Berlin? 



360 

Note. An extremely variable species, especially in the nature and 
degree of pubescence. Both pubescent and glabrous forms often grow 
together and, apparently, without ecological preference. Trautvetter, in 
describing his "var. setulosa," had in mind only peduncles ("pedicellis 
magis minusve glanduloso-setosis")- Actually, such or similar pubescence 
365 is also found at the base of the stem and quite often on the leaves. 
Branching varies less, being in the upper part of the stem, in the type 
specimen examined, but often beginning at the middle of the stem; 
occasionally the stem branches almost from the base. Sometimes other 
characters in this species (arrangement of leaves, degree of pubescence 
of the achenes, etc.) also vary. 

Section 2. Prenanthopsis Kirp. sect, nova in Addenda XXVIII, 726. — 
Mulgedium sect. II. Eumulgedium* cyanica Boiss. Fl. or III (1875) 800, 
p. p. — Achenes flat-trihedral or strongly compressed to flat, with numerous, 
more or less similar longitudinal ribs. 

Type of section: Cicerbita prenanthoides (M.B.) Beauverd. 

8. C. prenanthoides (MB.) Beauverd in Bull. Soc. Bot. Geneve, 2 
Ser. II (1910) 121 (seorsum impressum: Contrib. a I'etude des Compos. 
Ill, p. 23 No. 7); Grossh. Fl. Kavk. IV (1934) 252 and Opred. Rast. 
Kavt. 514; Kolak. Fl. Abkhazii IV, 303; Schchian in Fl. Gruzii, VIII, 
668. — Sonchus prenanthoides M.B. Fl. taur.-cauc. II (1808) 242. — 
Mulgedium prenanthoides DC. Prodr. VII, 1 (1838) 248; Ldb. Fl. Ross. 
II, 2, 842; Boiss. Fl. or. Ill, SOO. —Prenanthes erucaefolia Willd. ex Ldb. 
Fl. Ross. II, 2 (1845-1846) S42.—Lactuca prenanthoides Schmalh. Fl. II 
(1897) 167. — L. prenanthoides Bomm. in Mitt. Thiir. Bot. Ver. n. s. XX 
(1904-1905) 28 (non L. prenanthoides Scop. Fl. camiol. ed. 2, II (1772) 
100). — L. marschallii Stebbins in Joum. Bot. Lond. LXXV (1937) 16.— 
IMulgedium bourgaei Boiss. var. colchicum Alb. Prodr. fl. Colch. (1895) 
152 (Tr. Tifl. Bot. Sada, Prilozh. 1-e, Vyp. I).— 1 Cicerbita bourgaei var. 
p. colchica (Alb.) Beauverd, op. cit. p. 119. 

Perennial. Rhizome? Stem 80-120(200) cm high, shallow-sulcate, 
solid, rather densely covered with narrow, squamiform hairs, less often 
more or less arachnoid or almost glabrous. Leaves thin, elliptical 
(excluding uppermost), oblong-ovate or lanceolate, acute, short-cuspidate, 
cuneately narrowed at base or with winged petiole, with more or less 
prominent network of veins beneath, sometimes (especially along midrib) 
covered with narrow squamiform hairs or more or less glabrous, sinuate- 
toothed, with relatively long and narrow teeth; lower and middle leaves 
lyrate-lobate or lyrate, rarely lyrate or almost panduriform, with one or 
two pairs of lateral lobes and with large terminal, ovate or broadly 
triangular-ovate, lobe (rarely segment) often truncate at base; leaves on 



361 

inflorescence axis lanceolate or linear, small, those on peduncles reduced. 
Capitula mostly with 12-13-florets, almost cylindrical, with fruits 10- 
11(13) mm long, usually in narrowly racemose or paniculate-racemose 
inflorescence; peduncles slender, arachnoid-pubescent, with one to many 
366 scaly leaves. Involucral bracts three-rowed, dorsally finely appressed-hairy 
with papillose hairs or mixed with arachnoid hairs; inner bracts subobtuse, 
often with barbula of very short hairs (under a hand lens!). Corolla light- 
bluish. Achenes 4-5 mm long and about 1 mm wide, linear-ellipsoid or 
lanceolate, slightly narrowed toward both ends, somewhat flat or flat- 
trihedral, mature achenes dark violet, with numerous, prominent, more or 
less similar, longitudinal ribs, more or less densely covered with very 
short, upward-directed, stiff hairs (under high magnification!). Pappus two- 
rowed; hairs of outer row barely noticeable (under a hand lens!), dense, 
forming corona, those of inner row easily falling, thin, white, 5-7 mm 
long. Flowering (June) July to September. (Plate XXI, Fig. 8.) 

Montane forests and shrub thickets at altitudes from 800 to 2,000 
(3,000?) m. — Caucasus: Ciscaucasia, Eastern and Western Transcaucasia, 
endemic to the Caucasus, from where Marschall-Bieberstein described it. 
Type in Leningrad. 

Note. Cicerbita prenanthoides is close to C. bourgaei, with which it 
is sometimes confused. However, C. prenanthoides is distinguished from 
the latter by the leaf shape, much less developed inflorescence, and nature 
of pubescence on the disk bearing pappus. 

C. prenanthoides is a highly variable species, especially with respect 
to pubescence. 

9. C. bourgaei (Boiss.) Beauverd in Bull. Soc. Bot. Geneve, 2 Ser. 
II (1910) 118 (seorsum impressum: Contrib. a I'etude des Compos. Ill, p. 
20, No. 2); Grossh. H. Kavk. IV (1934) 252.— Mulgedium bourgaei Boiss. 
Fl. or. Ill (1875) 801. — ?A/. bourgaei Boiss. var. subintergrifolium Somm. 
and Lev. in Tr. Peterb, Bot. Sada XVI (1900) 297.— Exs: Balansa, PI. 
d'Or. 1866, No. 1467 (sub Mulgedio bourgaei Boiss.; isotypus); Sintenis, 
It. or. 1890, No. 3399 (sub eodem nomine). 

Perennial. Rhizome? Stem 80-150(300) cm high, shallow-sulcate, 
solid, densely covered with narrow, squamiform hairs in lower part, weakly 
pubescent or almost glabrous above. Leaves thin, but rather stiff, 
acuminate-elliptical or lanceolate (except uppermost), with distinct network 
of veins and scattered, narrow, squamiform hairs beneath, glabrous or 
subglabrous above, toothed or shallow sinuate-toothed, acute or cuspidate; 
lower leaves lyrate-lobate, usually with one pair of lateral lobes and large 
terminal lobe, narrowed toward base into broad-winged petiole with 
cordate-auriculate, semiamplexicaul base; middle cauline leaves sessile 
or subsessile, with base like lower leaves; upper leaves smaller and 



362 

narrower, often narrowed into petiole; usually linear-lanceolate, to linear 
in inflorescence, and scaly on inflorescence branches and peduncles. 
367 Capitula usually with 10-florets, almost cylindrical, with fruits 10-13 mm 
long, in more or less broadly pyramidal-paniculate, panculate or (less 
often) racemose-paniculate inflorescence, often large (30-50 cm long and 
more); inflorescence branches (usually 5-7 cm long, rarely shorter, but 
sometimes up to 30 cm long) terminating in slender, appressed arachnoid- 
pubescent, unequal peduncles. Involucral bracts two- or three-rowed; outer 
bracts with appressed-arachnoid and papillose pubescence on dorsal 
surface; inner ones with very short, papillose hairs (under high 
magnification!), subobtuse and often with beard of very short hairs. Corolla 
lilac-or azure-colored. Achenes about 5 mm long and slightly more than 
1 mm wide, linear-ellipsoid or lanceolate, sometimes slightly asymmetrical, 
somewhat flat, mature achenes dark violet, with numerous, strongly raised, 
almost similar, longitudinal ribs, rather densely covered with short, light- 
colored, upward-directed, stiff hairs (under a hand lens!), more or less 
distinctly narrowed at apex and bearing disk with pappus; disk densely 
covered with very fine hairs (under high magnification!). Pappus two- 
rowed; hairs of outer row very short and dense (under a hand lens!); 
those of inner row easily falling, thin, almost as long as achene. Flowering 
July to August. (Plate XXI, Fig. 9). 

Ravines and gullies, in shrub thickets, at edges and in logging areas 
of moist forests, and in tall-grass glades. — Caucasus: Ciscaucasia,? 
Dagestan (one doubtful specimen with the label of Lagovskii), ?Eastem 
Transcaucasia (western part). Western Transcaucasia (especially Abkhazia), 
General distribution: Balkans-Asia Minor (Turkey, Pontic Range). 
Described from Turkey. Type in Geneva; one isotype in Leningrad. 

Note. A Schchian (Zametka o Kavkhazskikh Vidakh roda Cicerbita 
[Note on the Caucasian species of Cicerbita]. Zam. po Sist. Geogr. Rast. 
Inst. Bot. Akad. Nauk GruzSSR, No. 13, 1947), believes that A. A. 
Grossheim carelessly reported C. bourgaei Boiss. from the Caucasus. 
According to Schchian, this species is "alien to our flora," and all the 
specimens identified by the Caucasian taxonomists as C. bourgaei should 
be referred to C. prenanthoides. However, the isotypes of C. bourgaei, 
preserved in the Herbarium of the Botanical Institute, Academy of Sciences 
of the USSR, allow me to resolve the question definitely in favor of this 
species occurring in the Caucasus (cf. also note on C prenanthoides). 

Section 3. Multicostaria Kirp. sect, nova in Addenda XXVIII, 726. — 
Achenes flat, often somewhat asymmetrical, with more or less numerous, 
somewhat similarly developed longitudinal ribs; achenes apically with 
more narrow, necklike constriction; outer row of pappus particularly well 
developed; corolla yellow (unlike other species of subgenus). 



363 

Type of section: Cicerbita deltoidea (M.B.) Beauverd. 

368 10. C. deltoidea (M.B.) Beauverd in Bull. Soc. Bot. Geneve, 2 Ser. 

II (1910) 128 (seorsum impressum: Contrib. a I'etude des Compos. Ill, p. 
30, No. 25) p. p. (solum quod epitheton Biebersteinii id est plantas 
georgicas attingit); Grossh. Fl. Kavk. IV (1934) 252 and Opred. Rast. 
Kavk. (1949) 514; Schchian in Fl. Gruzii VIII, 661 .—Prenanthes deltoidea 
M.B. Fl. taur.-cauc. Ill (1819) 539. — Mycelis ambigua Cass, in Diet. sc. 
nat. XL VIII (1827) A26.--Lactuca deltoidea C.A. Mey. Verzeichn. Pfl. 
Cauc. (1831) 56, p. p. (plantis talyschensibus exclusis); Ldb. Fl. Ross. II, 
2, 806, p. p.; Boiss. Fl. or. Ill, 816, p. p. — Cicerbita olgae Leskov in Tr. 
Bot. Muz. XXV (1932) 53; Grossh. op. cit (1934) 252 and (1949) 514; 
Kolak. in Fl. Abkhazii IV, 303; Schchian, op. cit. 668. — Ic: Leskov, op. 
cit. 54; Schchian, op. cit. 668, Plate 437. 

Perennial. Rhizome usually short. Stem (45)70-90 cm high, slightly 
sulcate, fistular, simple, glabrous or more or less densely covered with 
very narrow, squamiform, hairs below, weakly paniculately branched 
above, glabrous or subglabrous. Leaves thin, dark green above, gray or 
glaucous beneath, all or almost all leaves covered (especially on veins) 
with narrow, squamiform, hairs; lower leaves large, lyrately pinnatisect, 
with large hastate or ovate-hastate, irregularly toothed-spinulose terminal 
segment and smaller lateral segments of different form and size, but mostly 
oblong or lanceolate, slightly recurved; middle cauline leaves hastate, with 
broad-winged petiole and large amplexicaul auricles at base; leaves within 
inflorescence more or less strongly reduced. Capitula with 8-10-fIorets, 
cyUndrical or slightly expanded above, 10-12 mm long, borne on unequal 
peduncles in rather narrow and lax paniculate inflorescence. Involucre 
two- or three-rowed; inner involucral bracts usually violet at tip, with 
beard of very thin, light-colored hairs. Corolla yellow. Achenes about 4- 
5 mm long and 1 mm wide, narrow-ellipsoid, often asymmetrical, 
compressed, with large number (five to nine) of prominent ribs on both 
sides, mature achenes dark- or reddish-violet, covered with very short, 
light-colored hairs (under high magnification!), in sinuate rows; apically 
achenes distinctly attenuate into throat-like constriction terminating in a 
disk bearing pappus. Pappus two-rowed; outer row well developed, 
represented by dense corona of short, persistent, light-colored hairs; inner 
row of snow-white, fine, more or less easily falling, 5-6 mm-long hairs. 
Flowering June to August. (Plate XXII, Fig. 2). 

Middle mountain zone, in fir and beech-fir florests, less frequently 
in shrub thickets. — Caucasus: Ciscaucasia, Eastern Transcaucasia, 
?Westerm Transcaucasia. Endemic. Type in Leningrad. Described from 
Georgia. 

Note. A.I. Leskov, in describing his new species Cicerbita olgae. 



364 

noted: "Based on the structure of the florets, fruits, and capitula, our 
Cicerbita is almost indistinguishable from C. deltoidea (M.B.) Beauverd" 

369 (op. cit., p. 55). And, in fact, both these species are so similar that there 
is no real basis to treat C. olgae as a separate species. 

Subgenus III. Policilachena Kirp. subgen. nov. in Addenda XXVIII, 
726. — Achenes narrow or oblong-ellipsoid, ocherous or dark brown with 
dark violet spots, more or less distinctly compressed or not, with rather 
numerous, almost similarly developed longitudinal ribs and wholly covered 
with short hairs (under a hand lens!); achenes apically narrowed into 
short throat- like constriction terminating in broad disk bearing pappus. 
Pappus two-rowed; outer row sometimes indistinct, inner row of extremely 
fragile hairs. Capitula few-flowered (florets usually five, less often six 
and up to eight); corolla yellow or pink. Species growing in the mountains 
of Soviet Central Asia. 

Type of subgenus: Cicerbita zeravschanica M. Pop. ex S. Kovalevsk. 

11. C. zeravschanica M. Pop ex S. Kovalevsk. in Fl. Uzb, VI (1962) 
490, 517. — Mulgedium seravschanicum M. Pop. in Tr. Uzb. Gos. Univ. 
Nov. Ser. No. 27, Biol. Vyp. 14 (1941) 105 (descr. ross.).— Cicerbita 
seravschanica M. Pop. ex Zak. Fl. i Rastit. Bass. Zeravsh. II (1961) 427, 
nomen. — Ic- M. Pop. op. cit. (1941) fig. 3 (inter pages. 104 and 105). 

Perennial. Rhizome thicker than stem base, usually with dense tuft 
of numerous thin, funiform, roots. Stem 30-80(100) cm high, fistular, 
slightly sulcate, often (especially at base) reddish, glabrous or subglabrous 
below, more or less densely covered with short, squamiform, glandular 
hairs above. Leaves thin, green, sinuate or only slightly sinuate-toothed,, 
teeth spinescent, more or less whitish; lower leaves on long, narrow- 
winged, semiamplexicaul petioles, their laminas more or less densely 
covered with short hairs beneath, pinnatisect, terminating in hastate or 
cordate-ovate terminal segment and often also with two to many pairs of 
oblong, triangular, or irregular lateral segments reducing downward; leaves 
and petioles reduced toward stem apex, strongly expanded at base to 
auriculate-cordate and almost amplexicaul, laminas less divided (more or 
less entire in upper leaves); uppermost leaves oblong or lanceolate, scaly 
on peduncles. Capitula mostly with 5-florets, cylindrical, with fruits 7-10 
mm long, borne on slender, unequal peduncles, densely covered with 
short, squamiform, glandular hairs, in corymbose-paniculate or more or 
less corymbose, inflorescence. Involucral bracts usually two-rowed, green, 
dorsally densely covered with very short appressed pubescence (under a 
hand lens!) with more or less considerable mixture of erect glandular hairs. 
Corolla yellow. Achenes 4-5 mm long and about 1 mm wide, narrow 

370 or oblong-ellipsoid, slightly compressed, with rather numerous, prominent. 



365 

longitudinal ribs, ocherous or greenish-brown, with dark violet spots, 
somewhat densely covered throughout with short, light-colored, upward- 
directed hairs (under a hand lens!), more distinct in upper part; achenes 
apically attenuate into very short neck. Pappus two-rowed; outer row 
indistinct, inner one of fine, white, quite easily falling, about 5 mm-long 
hairs. Flowering June to September. (Plate XXII, Fig. 3.) 

Maple forests with grass-herb cover, juniper woodlands, rose thickets, 
herb-shrub thickets, slopes with shallow soil in thickets of Polygonum 
bucharicum, stony ravines, and alpine meadows (right up to glaciers?); at 
1,900-3,000 m. — Soviet Central Asia: Pamiro-Alai. Endemic. Described 
from Kugitang. Type in Tashkent. 

Note. An insufficiently studied and highly variable species. Short, 
eglandular plants with a weakly developed inflorescence are found together 
with (especially in Kylyab, Baldzhuan, Shuroabad and some other 
districts), tall plants (nearly 1 m high), with a strongly developed rhizome, 
relatively large inflorescence, and dorsally glandular involucral bracts. 
The material at my disposal does not allow me to attach taxonomic 
significance to all these differences. 

C. zeravschanica is close to C. rosea, but differs from it by the 
pubescence on the leaves beneath, yellow (and not pink) color of the 
corolla, pappus hairs not exserted beyond the involucre, smaller achenes 
without (as in C. rosea) a distinct throat-like constriction, as well as other 
characters. 

12. C. rosea (M. Pop. and Vved.) S. Kovalevsk. in H. Uzb. VI (1962) 
489. — Mulgedium roseum M. Pop and Vved, in Izv. Turkest. Otd. RGO 
XVII (1924) 29.— Ic: M. Pop. and Vved. op. cit. t. V (post page. 34); S. 
Kovalevsk. op. cit. t. XLIX, Fig. 1. 

Perennial. Rhizome rather long, almost as thick as stem base. Stem 
60-100 cm high, fistular, slightly sulcate, glabrous or almost glabrous 
from base to point of branching, densely covered above (especially from 
point of branching) with short, squamiform, glandular hairs. Leaves green 
above, more or less glaucous beneath, unevenly sinuate- 
372 toothed, with thick subulate tips; lower leaves on long, broad-winged, 
semiamplexicaul petiole, usually lyrately pinnatisect, with large hastate- 
or ovate-triangular terminal segment; toward stem apex leaves gradually 
reduced, petiole disappears, and lamina becomes entire, with cordate- 
auriculate, and almost amplexicaul base; uppermost leaves on inflorescence 
branches scaly. Capitula with five or six (eight)florets, cylindrical, with 
fruits 7-10 mm long, on unequal peduncles, densely covered with short 
glandular hairs, in corymbose-paniculate or more or less corymbose 
inflorescence. Involucral bracts mostly two-rowed, greenish, subobtuse 
and often with beard of thin white hairs, densely covered dorsally with 



366 




371 Plate XXIII. 

1 — Scariola orientalis (Boiss.) Sojak, habit of plant and achene; 2 — 5. viminea (L.) F.W. 

Schmidt, achene; 3 — Prenanthes purpurea L., achene; A — P. abietina (Boiss.) Kirp., achene; 

5 — P. tatarinowii Maxim., achene; 6 — P. cacaliifolia (M.B.) Beauverd, achene; 7 — P. 

pontica (Boiss.) Leskov, achene; 8 — P. angustifolia Boulos, achene; 9 — P. maximowiczii 

Kirp., achene. 



367 

very short, papillose hairs (under high magnification!). Corolla lilac- 
colored or whitish-pink. Achenes about 6 mm long and 1 mm wide, 
oblong-ellipsoid or lanceolate, slightly compressed, with many prominent 
longitudinal ribs, dark brown, covered with rather numerous, short, light 
colored, upward-directed hairs (under a hand lens!), more distinct in upper 
throat-like constriction; latter terminating in funnel-shaped disk bearing 
pappus; pappus hairs of outer row numerous, dense, white (under a hand 
lens!), inner row of very thin, white, very easily falling, about 5 mm- 
long hairs. Flowering May to August. (Plate XXII, Fig. 4). 

Stony and grassy slopes, in tall-grass cover, among shrubs and in 
spruce forests; at 1,000-2,400 m. — Soviet Central Asia: Pamiro-Alai 
(Uzgen, Gulcha), ?Syr-Darya, Tien Shan (mainly western part). Endemic. 
Described from western Tien Shan. Type in Tashkent. 

13. C. kovalevskiana Kirp. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk 
SSSR XXII (1963) 313. 

Perennial. Rhizome? Stem 100-110 cm high, fistular, sulcate, often 
(especially near base) reddish and rather thick (up to 10 mm), at base 
covered with individual squamiform hairs or glabrous, more or less 
glabrous above to point of branching, densely covered with squamiform 
glandular hairs in inflorescence. Leaves thin, green; lower leaves on long, 
broad-winged, almost amplexicaul petiole, more or less strongly 
pinnatisect, with large terminal segment; middle cauline leaves sessile, 
more or less elliptical or oblong-elliptical, with very broad, almost 
amplexicaul, cordate-auriculate or sagittate base, more or less distinctly 
sinuate-toothed, with spinescent teeth; leaves in inflorescence reduced. 
Capitula mostly with 5-florets, cylindrical, with fruits up to 1 cm long, 
on very slender peduncles much longer than capitula, densely covered 
373 with short, squamiform, glandular hairs; common inflorescence very long, 
usually very sparsely paniculate. Involucral bracts mostly two-rowed, 
green, dorsally covered with short, appressed, papillose hairs (under high 
magnification!), and often also with divergent glandular hairs; inner bracts 
longer, with membranous border. Corolla pink. Achenes about 4 mm long 
and 1 mm wide, oblong-ellipsoid or lanceolate, distinctly compressed, 
with numerous prominent longitudinal ribs, light brown, with scattered 
dark spots, densely covered throughout with short, light-colored upward- 
directed hairs, more distinct in upper part of achene; achenes apically 
attenuate into short but distinct, throat-like constriction; outer row of 
pappus comprising rather numerous, very short, white hairs (under a hand 
lens!), inner row of less numerous, fine, snow-white, more or less easily 
falling, about 5 mm long hairs. Flowering July to August. (Plate XXII, 
Figs. 1, 5). 

Deciduous forests and tree-shrub covered slopes. Soviet Central Asia: 



368 

Tien Shan (eastern Fergana). Endemic. Described from kyzyl-Ungur 
District. Type in Leningrad. 

Note. Originally, this species was designated by the name C. ferganica 
S. Kovalevsk. (cf. Kovalevskaja. liguliflorae flory Uzbekistana [Liguliflorae 
in the Flora of Uzbekistan]. Abstract of Dissertation. Tashkent. 1956, p. 
16) but was not described by her subsequently. 

Doubtful and Deleted Species 

1. C. oligocephala Zak. in Tr. Uzb. Gos. Univ., Nov. Ser. Vyp. 89, 
(1958) 166, nomen. 

It is reported from "Yailau" [summer pasture] and stony slopes with 
coarse herb cover. No more information reported. There is no description. 

2. C. plumieri (L.) Kirschleg. Fl. d' Alsace I (1852) 401.— Mulgedium 
plumieri C. Koch in Linnaea XXIII (1850) 668. 

Koch writes about tiiis plant: "Aus Grusien von Wilhelms erhalten." 
Meanwhile, there is no mention of Mulgedium plumieri by Boissier, nor 
by any of the authors of the flora of Caucasus, although Trautvetter 
included this species in his Incrementa. This is a western European plant, 
so the name given by Koch apparently is simply the result of an imprecise 
identification. 



GENUS 1660. Lagoseriopsis Kirp. Gen. Nov. 



1,2 



Kirp. in Addenda XXVIII, 726 

Capitula homogamous, few-flowered (with five or six florets), small, 
narrow-cylindrical or narrow-turbinate-cylindrical, solitary at apices of 
374